SOMATOSENSORY SYSTEM

External Environment through Touch

Position & Movement of our Body Parts (Proprioception) through Stimulation of Muscle & Joints
Temperature of the Body, Pain

2.1 Somatic Stimuli

Modality Specificity in the Somatosensory System.  The somatosensory systems process information about, and represent,
several modalities of somatic sensation (i.e., pain, temperature, touch, proprioception). Each of these modalities can be divided
into sub-modalities, as shown in Table 1 (e.g., pain into sharp, pricking, cutting pain; dull, burning pain; and deep aching
pain). Discriminative touch is also subdivided into touch, pressure, flutter and vibration. Each of these sensations (i.e., sub-
modalities) is represented by neurons that exhibit modality specificity. That is, when a somatosensory neuron is stimulated
naturally (e.g., by skin warming) or artificially (e.g., by electrical stimulation of the neuron), the sensation perceived is specific
to the information normally processed by the neuron (i.e., warm skin). Consequently, a "warm" somatosensory neuron will not
respond to cooling of the skin or to a touch stimulus that does not "warm" the skin. The somatosensory receptor and its central
connections determine the modality specificity of the neurons forming a somatosensory pathway.

Table I
The Sensory Modalities Represented by the Somatosensory Systems
Sub-Sub Somatosensory Pathway Somatosensory Pathway
Modality Sub Modality
Modality (Body) (Face)
sharp cutting pain   Neospinothalamic
Pain dull burning pain   Paleospinothalamic
deep aching pain   Archispinothalamic
Spinal Trigeminal
warm/hot   Paleospinothalamic
Temperature
cool/cold   Neospinothalamic
itch/tickle & crude touch   Paleospinothalamic
touch
Touch pressure
discriminative touch
flutter
vibration
muscle length
Position: Static Forces muscle tension Medial Lemniscal  Main Sensory Trigeminal
joint pressure
Proprioceptio
muscle length
n
Movement: Dynamic muscle tension
Forces joint pressure
joint angle

Tactile Stimuli. Tactile stimuli are external forces in physical contact with the skin that give rise to the sensations of touch,
pressure, flutter, or vibration. We normally think of touch as involving minimal force on-or-by an object that produces very
little distortion of the skin. In contrast, pressure involves a greater force that displaces the skin and underlying tissue. Time
varying tactile stimuli produce more complex sensations such as object movement or object flutter (20 to 50 Hz) or vibration
(100 to 300 Hz). An initial clinical examination of discriminative touch often involves testing the vibratory sense by applying a
128 Hz tuning fork over a bony prominence.

Proprioceptive Stimuli.1 Proprioceptive stimuli are internal forces that are generated by the position or movement of a body
part. Static forces on the joints, muscles and tendons, which maintain limb position against the force of gravity, indicate the
position of a limb. The movement of a limb is indicated by dynamic changes in the forces applied to muscles, tendons and
joints. An initial clinical examination of proprioception often involves testing the position sense by having the patient, with eyes
closed, touch one finger with another after the target finger has been moved. 

Proprioception is critical for maintaining posture and balance.  Somatosensory proprioceptive cues are combined with
vestibular proprioceptive cues and visual cues to control motor responses to changes in body/head position. During a clinical
examination, the Romberg test requires the patient to maintain balance while standing with feet together and eyes closed. It tests
whether the proprioceptive components are working properly when the visual cues are missing and proprioceptive cues are the
major sources of information. 

Sharp Cutting Pain Stimuli. Painful (nociceptive) stimuli are tissue-damaging sources of energy that may be external or
internal to the body surface. Sharp, cutting pain is the sensation elicited on initial contact with the painful stimulus. The
sensation of dull, burning pain may follow as a consequence of tissue inflammation. An initial clinical examination of the pain
sense often involves testing sharp, cutting pain sensitivity by asking the patient, who has her/his eyes closed, what they feel
when pricked with a pin. Pain mechanisms and pathways are described in detail in later chapters.

2.2 Introduction to Peripheral Organization of Somatosensory Systems

Peripheral Somatosensory Neurons. The cell bodies of the first-order (1°) somatosensory afferent neurons2 are located in
posterior root or cranial root ganglia (i.e., are part of the peripheral nervous system, Figure 2.1). The 1° afferents are
pseudounipolar cells. The cell body gives rise to a single process that divides to form a peripheral axon and a central axon. The
peripheral axon travels to and ends in the skin, muscle, tendon or joint and the central axon travels to and ends in the central
nervous system. 

Somatosensory Receptor Organ. The receptors of most sensory systems are located in specialized sensory receptor organs
(e.g., the photoreceptors in the eye and the auditory and vestibular hair cells in the inner ear) or within a restricted part of the
body (e.g., the taste buds in the mouth and the olfactory receptors in the olfactory mucosa of the nose). For the tactile
component of the somatosensory system, the skin covering the entire body, head and face functions as the touch receptor organ,
whereas joint tissues, muscles and tendons act as the proprioception receptor organs. These sensory receptor organs "house" the
somatosensory receptors and deliver the somatosensory stimuli to the receptors.

Sensory Receptors. Specialized sensory receptor cells (e.g., the photoreceptors of the eye) are located in specialized receptor
organs, produce receptor potentials, contain synaptic specializations, and release neural transmitters (Figure 2.2). Specialized
sensory receptors may be modified neurons (e.g., the photoreceptors and olfactory receptors) or modified epithelial cells (e.g.,
taste receptors and the auditory and vestibular hair cells).

 
Figure 2.1   Figure 2.2
The somatosensory first-order (1°) afferent is a pseudounipolar The specialized sensory receptors of the
neuron, which has a single process that divides into a peripheral auditory and visual systems. These cells
process and a central process. The peripheral process is part of the are specialized neurons (A. visual
peripheral nervous system (PNS) and terminates to form or end on a receptors) or specialized epithelial cells (B.
somatosensory receptor in skin, muscle or joint. The central process auditory receptors) that generate receptor
travels tso the central nervous system (CNS) where it terminates on potentials and contain synaptic vesicles.
a spinal cord or brain stem neuron.

There is only one type of sensory receptor cell in the somatosensory system, the Merkel cells, and they are found only in skin.
The vast majority of somatosensory receptors are not specialized receptor cells. That is, they are formed by the endings of the
somatosensory 1° afferent peripheral axon and adjacent tissue (Figure 2.3). There is no synaptic specialization or
neurotransmitter within the adjacent tissue. The adjacent tissue also does not generate receptor potentials.

  Figure 2.3
(A) When stimulated, the auditory receptor cell generates a receptor potential (1), which results
in the release of neurotransmitter at its synapse with the auditory 1° afferent. The
neurotransmitter depolarizes the 1° afferent, which generates action potentials (2 & 3) that travel
to the 1° afferent synaptic terminals on 2° afferents in the central nervous system. The 2°
afferent generates action potentials (4) in response to the transmitter release by the 1° afferent.
 (B) Most somatosensory receptors are not specialized receptor cells and are formed by the
terminal endings of the somatosensory 1° afferents. It is the 1° afferent terminal that produces a
  generator potential (1) which, in turn, initiates action potentials (2 & 3) in the 1° afferent axon.
The 1° afferent releases neurotransmitter on 2° afferents in the central nervous system. The 2°
afferent generates action potentials (4) in response to the transmitter by the 1° afferent.

Instead of ending on specialized receptors, most peripheral axons of somatosensory 1° afferents travel to skin, muscle or joint,
branch near their terminal sites, and end in the skin (Figure 2.4), muscle, tendon or joint tissue.

Figure 2.4
The primary
(1°)
somatosensor
y afferent
neuron. The
1° afferent's
cell body is
located in the
ganglion of a
cranial or
posterior
(spinal) nerve
root. The 1°
afferent's
peripheral
process
travels to
skin, muscle
or joint -
where it
branches into
terminal
fibers. Each
terminal fiber
forms, or ends
on, a
somatosensor
y receptor.
The 1°
afferent's
central
process joins
a cranial or
spinal nerve
and enters the
brain stem or
spinal cord -
where it
synapses with
a 2°
somatosensor
y neuron.

All the peripheral terminal branches of a 1° somatosensory axon end in a specific type of tissue (e.g., skin) and not in multiple
types of tissue (i.e., not in skin and muscle). All the peripheral terminal branches of a 1° axon form only one type of
somatosensory receptor. 

Figure 2.5
The locations
of
 somatosensor
y receptors in
the body.

Many of the 1° somatosensory afferent terminals are enveloped in a connective tissue capsule along with surrounding muscle,
tendon or cutaneous cells, or end on hair follicles. The hair follicles and the encapsulated tissue adjacent to the 1° afferent
terminals (i.e., skin, muscle, tendon, and joint tissues) contain no synaptic specializations and do not generate receptor
potentials or release neural transmitters. The complex of encapsulated tissue and afferent endings and the complex of hair
follicle and afferent endings play a role in the receptor transduction process, and each complex is considered to form a
"somatosensory receptor". Many other 1° somatosensory axons branch and terminate in skin, muscle, or joint as free nerve
endings. These endings are bare of myelin, are not encapsulated and are not associated with a specific type of tissue.

The sensitivity of the receptors to specific stimuli (e.g., touch verses muscle stretch) is determined by the location of the
receptor and by the non-neural tissue surrounding the 1° afferent terminal (Figure 2.6). 

Figure 2.6
The locations
of cutaneous
(somatosensor
y) receptors in
hairy and non-
hairy
(glabrous)
skin.

2.3 Sensory Transduction

The Adequate Stimulus. The adequate somatosensory stimulus (i.e., the stimulus to which a somatosensory neuron is most
sensitive) is either a mechanical force, a temperature change, tissue damage, or a chemical action. The discriminative touch and
proprioceptive systems are most sensitive to mechanical force. Consequently, their sensory receptors are of the
mechanoreceptor category.

Sensory Transduction. The non-neural tissue surrounding the peripheral ending of the somatosensory 1° afferent helps
concentrate and deliver the stimulus (e.g., mechanical force) onto the 1° afferent terminal membrane.
Somatosensory mechanoreceptors function to transduce the applied mechanical force into an electrical potential change in the
1° afferent neuron.

The mechanoreceptor 1° afferent terminal membrane contains ion channels that respond to mechanical distortion by increasing
sodium and potassium conductance (i.e., the channels are stress gated). Generator potentials are produced as sodium and
potassium flow down their electrochemical gradients to depolarize the terminal ending (see Figure 2.3B). In most cases, the
magnitude and duration of the generator potentials are related to the applied mechanical force: the greater the mechanical force,
the greater is the depolarization, and the longer the mechanical force is applied, the longer the terminal remains depolarized
(Figure 2.7). Terminals that do not sustain the depolarization for the duration of the mechanical distortion are called rapidly
adapting. Terminals that sustain the depolarization with minimal decrease in amplitude for the duration of a stimulus are called
slowly adapting.

Figure 2.7
At the TOP of
this figure,
two 1°
somatosensor
y neurons are
illustrated. A
mechanical
force (A) is
applied and
the responses
are measured
by a recording
electrode in
the
somatosensor
y receptor
(B), and a
recording
electrode in
the axon (C).
BELOW The
responses of
somatosensor
y 1° afferent
neurons to
stimulation of
the receptor
with a
sustained
stimulus are
illustrated for
rapidly
adapting
afferents
(LEFT panel)
and slowly
adapting
afferents
(RIGHT
panel). The
time course of
the applied
force or skin
displacement
(A); generator
potential
recorded in
the receptor
(B); and the
action
potentials
recorded from
the 1° afferent
axon (C) are
illustrated.
Notice that
the Ruffini
corpuscle and
Merkel disk
and their 1°
afferent
responses are
best suited to
transduce and
transmit
information
about long-
lasting
(maintained
or sustained)
stimuli that do
not vary over
time.
The generator potential spreads passively along the 1° terminal fiber to the axon trigger zone - that part of the 1° afferent axon
containing voltage-sensitive sodium and potassium channels (see Figure 2.3B). If the depolarization reaches threshold at these
voltage-sensitive sites, action potentials are generated by the 1° afferent peripheral axon. When the action potentials reach the
central terminals of the 1° afferent, they initiate the release neurotransmitters on 2° afferents within spinal cord or brain stem
nuclei. If, as in the example in Figure 2.8, the generator potential is slowly adapting, the 1° afferent produces a sustained
discharge of action potentials that continue for the duration of the stimulus.

Figure 2.8
Stretching the Ruffini corpuscle produces
a slowly adapting (sustained) generator
potential in the 1° afferent terminal that
degrades slowly for the duration of the
stretch. If the force applied to the 1°
afferent terminal produces a generator
potential that is of sufficient amplitude at
the axon trigger zone, a train of action
potentials is generated that travel along the
axon to the terminals of the its central
process. The action potentials in the
central terminals initiate the release of
neurotransmitters on 2° somatosensory
afferent neurons within the central
nervous system, which results in a
discharge of the 2° afferent.

If the generator potential is rapidly adapting (Figure 2.9), the 1° afferent produces a transient, short burst of action potentials
and falls silent even in the continued presence of the stimulus. 

Figure 2.9
Bending a hair produces a
rapidly adapting discharge
of action potentials in the 1°
afferent axon that does not
last the duration of the
bending force. If the force
applied to the 1° afferent
terminal produces a
generator potential that is of
sufficient amplitude at the
axon trigger zone, one or
more action potentials are
generated that travel to the
terminals of the 1° afferent
central process. The action
potentials in the central
terminals initiate the release
of neurotransmitters on 2°
somatosensory afferent
neurons within the central
nervous system. The 1°
afferent axon response is
rapidly adapting and action
potentials are only
generated when the hair is
bent.

The rapidly adapting receptors produce generator potentials and action potential discharges that follow the time-varying
waveform of pressure changes produced by a vibrating stimulus (Figure 2.10, left panel). In contrast, the slowing adapting
receptors produce generator potentials and action potential discharges that are sustained and unable to mimic the time-varying
pattern of the stimulus (Figure 2.10, right panel). Consequently, the responses of rapidly adapting 1° afferents are best suited for
representing time varying (e.g., vibrating or moving) stimuli, whereas slowly adapting 1° afferents better represent static stimuli
(e.g., sustained pressure).

Figure 2.10
At the TOP of this figure, two 1°
somatosensory neurons are
illustrated; each in contact with a
mechanical force (A), a recording
electrode in the somatosensory
receptor (B), and a recording
electrode in the axon (C). BELOW
The responses of the
somatosensory 1° afferents to
stimulation of the receptor with a
vibrating stimulus are illustrated
for rapidly adapting afferents
(LEFT panel) and slowly adapting
afferents (RIGHT panel). The time
course of the applied force or skin
displacement (A); generator
potential recorded in the receptor
(B); and the action potentials
recorded from the afferent axon are
illustrated (C). Notice that the
Pacinian and Meissner corpuscles
and their 1° afferent responses are
best suited to transduce and
transmit information about time-
varying (vibrating or moving)
mechanical stimuli.

2.4 Somatosensory Receptor Types

Figure 2.11
The locations of cutaneous receptors. Click on the somatosensory receptor
name (in green shaded area) to view a detailed drawing of the receptor. The
location of the receptor will be circled in the larger drawing of the skin.

Cutaneous Receptors

Some of the somatosensory receptors in skin (i.e., the cutaneous receptors) are classified as encapsulated receptors as the 1°
afferent terminal and surrounding cutaneous tissue are encapsulated by a thin sheath (Table II). The encapsulated cutaneous
receptors include Meissner corpuscles, Pacinian corpuscles and Ruffini corpuscles (See Figure 2.11). Other cutaneous receptors
are unencapsulated and include the hair follicle receptor (the 1° afferent ends on hair follicles) and the Merkel complex (the 1°
afferent ends at the base of a specialized receptor cell called the Merkel cell). The sensory receptors of the crude touch, pain and
temperature senses are bare or free nerve endings. That is, they are unencapsulated, do not end on or near specialized tissue, and
may be mechanoreceptors, nociceptors or thermoreceptors.

As was noted earlier, the sensitivity (modality specificity) of the somatosensory receptor is determined by its location and by the
structure of the non-neural tissue surrounding the 1° afferent terminal. The following describes the most commonly observed
cutaneous receptors.

Meissner Corpuscle. The Meissner corpuscle is found in glabrous (i.e., hairless) skin, within the dermal papillae (Figure
2.11). It consists of an elongated, encapsulated stack of flattened epithelial (laminar) cells with 1° afferent terminal fibers
interdigitated between the cells (Figure 2.12).

  Figure 2.12
 The Meissner
corpuscle consists of
an encapsulated stack
of flattened epithelial
(laminar) cells with 1°
afferent terminals
interdigitated between
these cells. The
Meissner corpuscle is
located within the
dermal papilla, near
the surface of the skin,
with its long axis
perpendicular to the
skin surface.

A force applied to non-hairy skin (Figure 2.13) causes the laminar cells in the Meissner corpuscle to slide past one another,
which distorts the membranes of the axon terminals located between these cells. If the force is maintained, the laminar cells
remain in a fixed, albeit, displaced position, and the shearing force on the axon terminals' membranes disappears. Consequently,
the 1° afferent axons produce a transient, rapidly adapting response to a sustained mechanical stimulus.

Figure 2.13
When a force is applied to the dermal papilla containing the Meissner
corpuscle, the laminar cells in the corpuscle slide past one another. This
shearing force distorts the membranes of the axon terminals located between
the laminar cells, which depolarizes the axon terminals. If the force is
sustained on the dermal papilla, the laminar cells remain in their displaced
positions and no longer produce a shearing force on the axon terminals.
Consequently, a sustained force on the dermal papilla is transformed into a
transient force on the axon terminals of the Meissner corpuscle. The 1°
afferent axon response of a Meissner corpuscle is rapidly adapting and action
potentials are only generated when the force is first applied.

The Meissner 1° afferent discharges "follow" low frequency vibrating (30 -50 Hz) stimuli, which produces the sensation of
"flutter" (Figure 2.10, left panel). Because a single 1° afferent axon forms many, dispersed (3-4 mm) Meissner corpuscles, the
1° afferent can detect and signal small movements across the skin. Stimulation of a sequence of Meissner corpuscles have been
described to produce the perception of localized movement along the skin.

Consequently, Meissner corpuscles are considered to be the discriminative touch system's flutter and movement detecting
receptors in non-hairy skin.

Pacinian Corpuscle. Pacinian corpuscles are found in subcutaneous tissue beneath the dermis (Figure 2.9) and in the
connectivetissues of bone, the body wall and body cavity. Therefore, they can be cutaneous, proprioceptive or visceral
receptors, depending on their location.

Figure 2.14
The Pacinian corpuscle consists of a single, centrally placed 1° afferent
terminal that is surrounded by concentrically layered epithelial (laminar) cells
that are all encapsulated within a sheath. In skin, the Pacinian corpuscle is
located deep in the subcutaneous adipose tissue.

The Pacinian corpuscle is football-shaped, encapsulated, and contains concentrically layered epithelial (laminar) cells (Figure
2.14). In cross section, the Pacinian corpuscle looks like a slice of onion, with a single 1° afferent terminal fiber located in its
center. The outer layers of laminar cells contain fluid that is displaced when a force is applied on the corpuscle.

When a force is first applied on the Pacinian corpuscle (Figure 2.15), it initially displaces the laminar cells and distorts the axon
terminal membrane. If the external pressure is maintained on the corpuscle, the displacement of fluid in the outer laminar cells
dissipates the applied force on the axon terminal. Consequently, a sustained force on the corpuscle is transformed into a
transient force on the axon terminal, and the Pacinian corpuscle 1° afferent produces a fast adapting response.
 Figure 2.15
When a force is applied to
the tissue overlying the
Pacinian corpuscle (press
PLAY), its outer laminar
cells, which contain fluid,
are displaced and distort
the axon terminal
membrane. If the pressure
is sustained on the
corpuscle, the fluid is
displaced, which dissipates
the applied force on the
axon terminal.
Consequently, a sustained
force on the Pacinian
corpuscle is transformed
into a transient force on its
axon terminal. The
Pacinian corpuscle 1°
afferent axon response is
rapidly adapting and
action potentials are only
generated when the force
is first applied.

Pacinian corpuscles 1° afferent axons are most sensitive to vibrating stimuli (e.g., a tuning fork vibrating at 100 to 300 Hz,
Figure 2.10, left) and unresponsive to steady pressure. The sensation elicited when cutaneous Pacinian corpuscles are stimulated
is of vibration or tickle.

Pacinian corpuscles in skin are considered to be the vibration sensitive receptors of the discriminative touch system.

Ruffini Corpuscle. The Ruffini corpuscles are found deep in the skin (Figure 2.11), as well as in joint ligaments and joint
capsules and can function as cutaneous or proprioceptive receptors depending on their location. The Ruffini corpuscle (Figure
2.16) is cigar-shaped, encapsulated, and contains longitudinal strands of collagenous fibers that are continuous with the
connective tissue of the skin or joint. Within the capsule, the 1° afferent fiber branches repeatedly and its branches are
intertwined with the encapsulated collagenous fibers. 

Figure 2.16
The Ruffini corpuscle consists of 1°
afferent terminal fibers that are
intertwined with collagenous fibers
and together with the collagenous
fibers are encapsulated in a fibrous
sheath. The Ruffini corpuscles are
oriented parallel to the skin surface
and situated deep within the dermis.

The Ruffini corpuscles are oriented with their long axes parallel to the surface of the skin and are most sensitive to skin stretch.
Stretching the skin (Figure 2.17) stretches the collagen fibers within the Ruffini corpuscle, which compresses the axon
terminals. As the collagen fibers remain stretched and the axon terminals remain compressed during the skin stretch, the Ruffini
corpuscle's 1° afferent axon produces a sustained slowly adapting discharge to maintained stimuli.

Figure 2.17
When the skin is stretched, the
collagen fibers in the Ruffini
corpuscles are also stretched and
compress their 1° afferent
terminals. As the collagen fibers
 remain stretched and the axon
terminals remain compressed
during the skin stretch, the
Ruffini corpuscle 1° afferent
axon produces a sustained
generator potential and a slowly
adapting discharge to
maintained stimuli.

Ruffini corpuscles in skin are considered to be skin stretch sensitive receptors of the discriminative touch system. They also
work with the proprioceptors in joints and muscles to indicate the position and movement of body parts.

Hair Follicle. The hair follicle receptor is an unencapsulated cutaneous receptor (Figure 2.10). The 1° afferent terminal
axons spiral around the hair follicle base or run parallel to the hair shaft forming a lattice-like pattern (Figure 2.18). 

Figure 2.18
The hair follicle 1°
afferent terminal
fibers enter the
follicle to encircle or
to form a lattice
pattern around the
hair shaft.

Most hair follicle 1° afferents are the fast-adapting type; displacement of the hair produces a transient discharge of action
potentials at the onset of the displacement and a maintained displacement of the hair often fails to produce a sustained discharge
(Figure 2.19). The hair follicle afferents respond best to moving objects and signal the direction and velocity of the movement
of a stimulus brushing against hairy skin.

Figure 2.19
Bending a hair produces a
transient force on the hair
follicle base as the entire
follicle is displaced by the
bending force. The 1°
afferent terminal may
produce a rapidly adapting
generator potential and the
1° afferent axon a transient
discharge of action
potentials — even to
sustained bending of the
hair.

As Meissner corpuscles are absent from hairy skin, the hair follicle endings are considered to be the discriminative touch
system's movement sensitive receptors in hairy skin.

Merkel Complex. The Merkel complex is found in both hairy and non-hairy skin and is located in the basal layer of the
epidermis (Figure 2.11). The Merkel complex is unencapsulated and consists of a specialized receptor cell, the Merkel cell, and
a 1° afferent terminal ending, the Merkel disk3 (Figure 2.20). Thick, short, finger-like protrusions of the Merkel cell couple it
tightly to the surrounding tissue. The Merkel cell is a modified epithelial cell, which contains synaptic vesicles that appear to
release neuropeptides that modulate the activity of the 1° afferent terminal. Each 1° afferent axon often innervates only a few
Merkel cells in a discrete patch of skin (Figure 2.18). 

Figure 2.20
The Merkel complex consists of a
specialized Merkel cell, which
 contains synaptic vesicles, and the
Merkel disk ending of a 1° afferent
terminal fiber. A single 1° afferent
axon often innervates only a few
Merkel cells within a discrete patch of
skin.

A force applied to the skin overlying the Merkel cell distorts it (Figure 2.21), which stimulates its release of a neuropeptide at its
synaptic junctions with the Merkel disk. As the Merkel cell is mechanically coupled to the surrounding skin, it remains distorted
for the duration of the force applied on the overlying skin. Consequently, the Merkel complex 1° afferent axon responds to
small forces applied to a discrete patch of skin with a slowly adapting, sustained discharge.

Figure 2.21
The Merkel cell is
coupled to the
surrounding tissue and
cannot shift its
position relative to the
surrounding tissue.
Consequently, a force
applied to the
overlying skin
(press PLAY), distorts
the Merkel cell for the
duration of the applied
force. The distortion
of the Merkel cell
results in the release of
a stream of
neuropeptides at its
synaptic junctions
with the 1° Merkel
disk. As a result the
action potential
discharges produced
by the Merkel
complex 1° afferent is
slowly adapting.

Merkel cells are considered to be the fine tactile receptors of the discriminative touch system that provide cues used to localize
tactile stimuli and to perceive the edges (shape or form) of objects.

Free Nerve Endings. Free nerve endings are found throughout the body, in skin (Figure 2.11), muscles, tendons, joints,
mucous membranes, cornea, body mesentery, the dura, the viscera, etc. The free nerve endings in skin are stimulated by tissue-
damaging (nociceptive) stimuli that produce the sensation of pain or by cooling of the skin or the warming of skin or by touch.
Notice that although all cutaneous free nerve endings appear very similar morphologically, there are different functional types
of free nerve endings, with each responding to specific types of cutaneous stimuli (e.g., nociceptive, cooling, warming or
touch).

Free nerve endings are considered to be the somatosensory receptors for pain, temperature and crude touch.

Table II
Cutaneous Receptors
Receptor Type Sensation Signals Adaptation
Meissner  Encapsulated  Touch: Flutter &
Frequency/Velocity & Direction  Rapid
corpuscle  & layered  Movement
Pacinian  Encapsulated 
Touch: Vibration  Frequency: 100-300 Hz  Rapid
corpuscle  & layered 
 Ruffini  Encapsulated 
Touch: Skin Stretch  Direction & Force Slow
corpuscle  collagen 
Direction & 
Hair follicle Unencapsulated  Touch: Movement  Rapid
Velocity
Merkel  Specialized 
Touch, Pressure, Form  Location & Magnitude Slow
complex  epithelial cell 
Free Nerve  Pain, Touch, or Tissue damage, Contact, or Depends on information
Unencapsulated 
Ending  Temperature  Temperature change carried

2.5 Proprioceptive Receptors 
Proprioceptors are located in muscles, tendons, joint ligaments and in joint capsules. There are no specialized sensory receptor
cells for body proprioception4. In skeletal (striated) muscle, there are two types of encapsulated proprioceptors, muscle spindles
and Golgi tendon organs (Figure 2.22), as well as numerous free nerve endings. Within the joints, there are encapsulated
endings similar to those in skin, as well as numerous free nerve endings.

Figure 2.22
A muscle spindle
receptor and Golgi
tendon organ in the
bicep muscle.

Muscle Spindles. Muscle spindles are found in nearly all striated muscles. A muscle spindle is encapsulated and consists of
small muscle fibers, called intrafusal muscle fibers, and afferent and efferent nerve terminals (Figure 2.23).

Figure 2.23
A muscle spindle with its
sensory and motor
innervation. The primary
muscle spindle afferent
terminates as annulospiral
endings in the central area
of the intrafusal muscles
whereas the secondary
muscle spindle afferent
terminates as flower spray
endings in more polar
regions of intrafusal
muscles. The motor
endplates of gamma motor
neurons are located in the
polar regions. The muscle
spindle is attached to the
surrounding extrastriate
muscles and lays with its
long axis in parallel with
the long axes of the
surrounding muscle. 

Intrafusal muscles are found exclusively in muscle spindle receptors and are distributed throughout the body among the
ordinary extrafusal muscle fibers of skeletal muscles. The intrafusal fibers are attached to the larger, surrounding extrafusal
muscle fibers. They are oriented in parallel with the extrafusal fibers but do not contribute directly to muscle strength when they
contract because of their small size. 

There are two types of afferent terminals in the muscle spindle (Figure 2.23). The annulospiral endings wrap around the central
region of the intrafusal fibers, whereas the flower-spray endings terminate predominantly in more polar regions (away from the
central area) of the intrafusal fibers. The 1° afferents forming the annulospiral endings are called the primary muscle spindle
afferents, whereas those forming the flower-spray endings are called the secondary muscle spindle afferents.
In addition to afferent terminals, the terminals (motor endplates) of gamma motor neurons end on intrafusal muscle fibers. They
will be described in detail in the chapters covering motor systems.

In summary, the muscle spindles are proprioceptors specialized to monitor muscle length (stretch) and signal the rate of change
in muscle length by changing the discharge rate of afferent action potentials. Muscle spindles are most numerous in muscles that
carry out fine movements, such as the extraocular muscles and the intrinsic muscles of the hand. There are fewer spindles in
large muscles that control gross movements of the body (e.g., the muscles of the back). 

Figure 2.24
The Golgi tendon
organ is located
at the junction of
muscle and
tendon. The
Golgi tendon
organs resemble
the Ruffini
corpuscles. That
is, the 1° afferent
terminal fibers
are intertwined
with collagenous
fibers of the
tendon and the
entire organ is
encapsulated in a
fibrous sheath.

Golgi Tendon Organs. Golgi tendon organs are found in the tendons of striated extrafusal muscles near the muscle-tendon
junction (Figure 2.22). Golgi tendon organs resemble Ruffini corpuscles. For example, they are encapsulated and contain
intertwining collagen bundles, which are continuous with the muscle tendon, and fine branches of afferent fibers that weave
between the collagen bundles (Figure 2.24). They are functionally "in series" with striated muscle.

The Golgi tendon organ collagen fibers are continuous with the extrafusal muscle at one end and with the muscle tendon at its
opposite end. Consequently, the mechanical force on the organ is maximal when the extrafusal muscles contract, shorten, and
increase the tension on the tendon. When the muscles contract, the 1° afferent terminals are compressed and remain compressed
as long as the muscle remains contracted. The Golgi tendon organ 1° afferent response to sustained isometric muscle
contraction is slowly adapting, and the 1° afferent generates action potentials as long as the tension is maintained. The responses
of the Golgi tendon organ 1° afferent axon is maximal when the contracted muscle bears a load, e.g., when lifting a heavy
object.

The Golgi tendon organ  is a proprioceptor that monitors and signals muscle contraction against a force (muscle tension),
whereas the muscle spindle is a proprioceptor that monitors and signals muscle stretch (muscle length). 

Joint Receptors. Joint receptors are found within the connective tissue, capsule and ligaments of joints (Figure 2.25). The
encapsulated endings resemble the Ruffini and Pacinian corpuscles and the Golgi tendon organs.

Figure 2.25
The joint receptors are
free nerve endings and
encapsulated endings
in the joint capsule and
joint ligaments. The
encapsulated receptors
in the joint capsule
resemble Pacinian and
Ruffini endings
whereas those in the
ligaments resemble
 Golgi tendon organs.

The joint 1° afferents respond to changes in the angle, direction, and velocity of movement in a joint. The responses are
predominantly rapidly adapting with few joint 1° afferents signaling the resting (static) position of the joint. It has been
suggested that information from muscles, tendons, skin and joints are combined to provide estimates of joint position and
movement. For example, when the hip joint is replaced — removing all joint receptors — the ability to detect the position of the
thigh relative to the pelvis is not lost. 

Free Nerve Endings. As mentioned above, free nerve endings of 1° afferents are abundant in muscles, tendons, joints, and
ligaments. These free nerve endings are considered to be the somatosensory receptors for pain resulting from muscle, tendon,
joint, or ligament damage and are not considered to be part of the proprioceptive system.

Table III
Receptor Type Sensation Signals Adaptation
Encapsulated
Rapid initial
annulospiral and Muscle   Muscle 
Muscle Spindle  transient and slow
flower spray  stretch  length & velocity 
sustained
endings 
Muscle: Golgi  Encapsulated  Muscle 
Muscle tension  Slow
Tendon Organ  collagen  contraction
Joint:  Encapsulated  Direction &
Joint Movement  Rapid
Pacinian  & layered  velocity
Joint:  Encapsulated 
Joint pressure  Pressure & Angle Slow
Ruffini  collagen 
Joint: Golgi  Encapsulated 
Joint torque  Twisting force Slow
Organ  collagen 

2.6 Summary
In this chapter, you have learned about somatosensory stimuli and the receptors of three components of the somatosensory
systems. These three components provide accurate information about the location, shape, texture, and movement of tactile
stimuli, (discriminative touch), the position and movement of body parts (proprioception) and the application and location of
painful stimuli (nociception). Tactile and proprioceptive stimuli are the mechanical forces produced when skin contacts external
objects (discriminative touch), limbs oppose the force of gravity (body position) and muscles contract and body parts move.
Painful stimuli are tissue-damaging forces. The sensations produced are those of touch, pressure, flutter, and
vibration/movement (discriminative touch), body position and movement (proprioception), and sharp cutting pain. The
discriminative touch receptors are encapsulated 1° afferent terminals (Meissner, Pacinian and Ruffini corpuscles), hair follicle
endings and Merkel complexes in skin. The proprioceptive receptors in muscle are also encapsulated and include the muscle
spindle and Golgi tendon organ. The joint receptors are similar to the encapsulated endings in skin and tendon and are found in
the joint capsule and ligaments. The sharp cutting nociceptors are free nerve endings. 

Although it is convenient to subdivide somatosensory receptors and pathways for didactic, clinical and research purposes, it is
important to keep in mind that most somatosensory stimuli act simultaneously and in varying degrees on all somatosensory
receptors in the body part stimulated. For example, placing a heavy, cold object in an outstretched hand produces tactile,
thermal, and proprioceptive sensations that allow us to appreciate the presence (touch, pressure), temperature, and weight of the
object and provide proprioceptive information for finger, wrist and arm adjustments so we do not drop the object