Chapter 2: Somatosensory Systems

Patrick Dougherty, Ph.D., Department of Anesthesiology and Pain Medicine, MD Anderson

Cancer Center
(content provided by Chieyeko Tsuchitani, Ph.D.)

The somatosensory systems inform us about objects in our external environment

through touch (i.e., physical contact with skin) and about the position and
movement of our body parts (proprioception) through the stimulation of muscle and
joints. The somatosensory systems also monitor the temperature of the body,
external objects and environment, and provide information about painful, itchy and
tickling stimuli. The sensory information processed by the somatosensory systems
travels along different anatomical pathways depending on the information carried.
For example, the posterior column-medial lemniscal pathway carries discriminative
touch and proprioceptive information from the body, and the main sensory
trigeminal pathway carries this information from the face. Whereas, the
spinothalamic pathways carry crude touch, pain and temperature information from
the body, and the spinal trigeminal pathway carries this information from the face.
This first series of chapters on somatosensory systems concentrates on the
somatosensory systems that provide accurate information about the location and
temporal features of stimuli and about sharp pain, tactile stimuli and the position
and movement of body parts. This chapter describes somatosensory stimuli, the
sensations produced when they are applied, and the cutaneous, muscle, and joint
receptors that are responsible for initiating the perceived somatic sensations.
Subsequent chapters describe the pathways processing other pain, temperature,
crude touch and visceral sensations.

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 Sy

Modality
Pain
Temperature

Touch

Sub Modality
sharp cutting pain
dull burning pain
deep aching pain
warm/hot
cool/cold
itch/tickle & crude touch
discriminative touch

Position: Static Forces

Proprioception
Movement: Dynamic Forces

Sub-Sub Modality

touch
pressure
flutter
vibration
muscle length
muscle tension
joint pressure
muscle length
muscle tension
joint pressure
joint angle

Somatosensory Pathway (Body)

Neospinothalamic
Paleospinothalamic
Archispinothalamic
Paleospinothalamic
Neospinothalamic
Paleospinothalamic

Medial Lemniscal

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
The specialized sensory receptors of
pseudounipolar neuron, which has a single process that
the auditory and visual systems.
divides into a peripheral process and a central process. The These cells are specialized neurons (A.
peripheral process is part of the peripheral nervous system
visual receptors) or specialized
(PNS) and terminates to form or end on a somatosensory
epithelial cells (B. auditory receptors)
receptor in skin, muscle or joint. The central process travels that generate receptor potentials and
tso the central nervous system (CNS) where it terminates on
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)
somatosens
ory 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
somatosens
ory 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
somatosens
ory 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
somatosens
ory
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
(somatosens
ory)
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.
Somatosensorymechanoreceptors 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
somatosens

ory neurons
are
illustrated.
A
mechanical
force (A) is
applied and
the
responses
are
measured by
a recording
electrode in
the
somatosens
ory receptor
(B), and a
recording
electrode in
the axon (C).
BELOW The
responses of
somatosens
ory 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
displacemen
t (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 longlasting
(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 disk 3 (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
(pressPLAY),
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.

Receptor
Meissner
corpuscle

Type
Encapsulated
& layered

Table II
Cutaneous Receptors
Sensation

Touch: Flutter & Movement

Signals

Frequency/Velocity & Direction

Pacinian
corpuscle
Ruffini
corpuscle

Encapsulated
& layered
Encapsulated
collagen

Hair follicle
Merkel
complex
Free Nerve
Ending

Touch: Vibration

Frequency: 100-300 Hz

Touch: Skin Stretch

Direction & Force

Unencapsulated

Touch: Movement

Direction &
Velocity

Specialized
epithelial cell

Touch, Pressure, Form

Location & Magnitude

Unencapsulated

Pain, Touch, or Temperature

Tissue damage, Contact, or Tempera

change

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 proprioception 4. 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

Muscle
Spindle

Encapsulated
annulospiral
and flower
spray
endings

Muscle
stretch

Muscle
length &
velocity

Rapid initial
transient and
slow
sustained

Muscle:
Golgi
Tendon
Organ

Encapsulated
collagen

Muscle
tension

Muscle
contraction

Slow

Joint:
Pacinian

Encapsulated
& layered

Joint
Movement

Direction &
velocity

Rapid

Joint:

Encapsulated

Joint

Pressure &

Slow

Ruffini

collagen

pressure

Angle

Joint:
Golgi
Organ

Encapsulated
collagen

Joint torque

Twisting
force

Slow

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.

Somatosensation
Written by: Andrea Alenda from UCL, Greta Santagata from Manchester University,

Introduction to the Somatosensory System

How many senses do humans have? Five? You have known this since you were a kid, but, after
reading this article, you might want to reconsider.
The somatosensory system detects, relays and processes somatosensory sensations, like touch,
vibration, pressure, itch, nociception (information about painful stimuli), temperature, proprioception
(information about the position and movement of our joints and muscles) and interoception
(information about internal organs). So, without taking into account what other sensory systems do,
the somatosensory system is already detecting more than five senses by itself.

The somatosensory system has been linked to the sense of self awareness as it provides an internal
representation of the body. In pathologies affecting the somatosensory system, such as the phantom
limb or the hemineglect syndrome, this awareness of the body is challenged.
This article will look at the basic anatomy and physiology of the somatosensory system, from
receptors in the periphery to the cerebral cortex in the brain.

Characteristics of somatosensory receptors

To explain the properties of the somatosensory receptors we are going to use two important concepts:

Receptive field: the physical space where a stimulus has to

appear in order to be detected by the receptor. This physical
space can have different sizes.

Adaptation is a physiological property of the neural response to

stimuli measured in the number and timing of action potentials.
Commonly a neuron that shows adaptation will respond with a
large number of action potentials when being stimulated but will
gradually decrease the number of action potentials over time.
This is because it quite literally gets "used" to the stimulus after
a while. Each type of mechanoreceptor can be either rapidly or
slowly adapting (RA/SA). The rapidly adapting
mechanoreceptors fire when they detect a change in their
receptive field. For example they can get activated when a
stimulus is delivered or withdrawn, but they won't fire during the
period of time the stimulus is present in the receptive field.
Slowly adapting mechanoreceptors will respond throughout the
whole duration of the stimulus in the receptive field.

Different receptors for different modalities

The somatosensory system is generally considered to have four
modalities:mechanoreception, propioception,temperature and nociception. Each of these
modalities has its specialised receptors, which will transmit their information through specific
anatomical pathways to the brain.

The mechanoreceptors
Of the four modalities we will concentrate on touch.
Mechanoreceptors deal with mechanical stimuli, such as pressure and vibration. Most
mechanoreceptors are found in the skin, although some are also found in the muscles, tendons and
joints.
There are four types of skin mechanoreceptors:

Meissner's corpuscules convey information regarding stroking

and fluttering. They are characterized by a small receptive field
and a rapid adapating response (RA).

Merkel's disks convey information regarding pressure and

texture. It is characterized by a small receptive field and a slow
adapting response (SA)

Ruffini's capsules respond to stretching of the skin. They have a

big receptive field and and have the slowest adapting response of
all mechanoreceptors (SA)

Pacinian corpuscules: respond to vibration. Their receptive field

is large and they show an extremely rapid adapting response (RA)

Speed of information travelling to the brain

Anatomical pathways are composed by populations of neurons. The speed by which information will
reach the brain will depend on the type neurons that compose the anatomical pathway. To be more
precise, by the size and myelination of the axons of these neurons. The bigger and the more
myelinated* the axon is, the faster the signal will travel. Depending on these characteristics, axons are
classified as:

A alpha: biggest diameter and fully insulated by the myelin

sheath. These fibres are the quickest to transmit information.
Proprioceptores of skeletal muscle are innervated by this type of
fibre.

A beta: medium size diameter and myelined. Mechanoreceptors

in the skin are innervated by this type of fibre.

A delta: small diameter with little myelination. They relay

temperature and information on pain.

C : small diameter and non-myelinated fibres, they relay

information on temperature, pain and itch.

* Myelin is a sheath that covers the axons of some neurons. The more myelinated an axon is, the
faster the signal will travel through it, because the myelin sheath increases the speed of the electrical
signal travelling through the axon by decreasing capacitance and increasing electrical resistance.
Experiment yourself
You fall on a pool (with water) belly first. What will you experience? First you will perceive the position
of your body followed by the temperature of the water. Then you will have a quick acute pain in the
stomach, the first area that contacted the water. Finally, a pain that starts a few miliseconds later,
which makes your body curl. What happened? The A alpha fibers have transmitted information about
the position of your body faster, then the A delta transmit the information of temperature and acute
pain. Finally information carried through the C fibers reached the brain with the slow and more intense
pain.

The somatosensory pathways

Axons from sensory receptors are segregated into parallel anatomical pathways that reach different
processing areas in the brain. Each class of sensory receptor is capable of activating a corresponding
cortical area. This is known asmodality segregation, which allows the brain to reconstruct signals by
time, modality and location. This is true for all sensory systems.
Modality specific information travels from the receptors to the brain through different anatomical
pahtways:

Two somatosensory pathways for the body: the anterolateral

pathway and the dorsal column/medial lemnical pathway

Two somatosensory pathways for the head: the spinotrigeminal

pathway and the trigeminal pathway.

The ascending pathways: from somatosensory receptor to

cerebral cortex
This is a summary of the ascending pathways carrying somatosensory information to the brain, the
descending pathways will not be covered here.
Information from the body:

Discriminative touch - the dorsal column/medial lemniscal

system

Large-diameter afferents (A-alpha and A-beta fibres) synapse in the dorsal column nuclei of the
medulla (gracile and cuneate), then cross the midline and ascend to the ventro posterolateral (VPL)
thalamic nucleus, via the medial lemniscus. They then finally reach the somatosensory cortex.
Information of discriminative touch and proprioception travels through fast conducting fibres. The
afferents that compose this pathway cross the midline, which means that the right hemisphere of the
brain will process the information regarding the left side of the body and viceversa.

Nociception and temperature the anterolateral system

Smaller afferents (A-delta and C fibres) synapse in the spinal cord, then cross midline and ascend via
the spinal cord and brainstem to the VPL and other nuclei of the thalamus. Collaterals of these axons
terminate in the reticular formation of the pons and medulla. Information about temperature, deep
touch and nociception travels through slow conducting fibres. The anterolateral system is complex, it
splits into three pathways, each of which ends in different brain areas with several relay synapses.
Awareness of pain, temperature and deep touch is distributed through different brain areas. The
afferents that compose this pathway also cross the midline, this means that the right hemisphere of
the brain will process the information regarding the left side of the body and viceversa.
Information from the head:

Discriminative touch - the trigeminal system

Large, fast axons (A-alpha and A-beta) innervate the pars oralis and principal sensory nucleus. The
second order axons cross the midline, ascend in the trigeminothalamic tract and terminate in the
ventro posteromedial nucleus (VPM). From the thalamus information is conducted to the
somatosensory cortex. The afferents that compose this pathway cross the midline, which means that
the right hemisphere of the brain will process the information regarding the left side of the head and
viceversa.

Nociception and temperature - the spinotrigeminal system

Small, slowly conducting axons (A-delta and C fibres) descend in the spinal trigeminal tract and
terminate in the pars caudalis of the spinal nucleus of the brainstem. The second-order axons cross
the midline and ascend to the VPM and intralaminar nuclei of the thalamus. From the thalamus
information reaches the somatosensory cortex. The afferents that compose this pathway cross the
midline, which means that the right hemisphere of the brain will process the information regarding the
left side of the head and viceversa.

The dorsal column/medial lemniscal pathway

The anterolateral system

The trigeminal pathway

The Spinotrigeminal pathway

The thalamus
The thalamus is a structure with the shape of two attached beetles that belongs to the diencephalon. It
is essential for gating, processing and transferring information to and from the cerebral cortex. The
majority of sensory information coming from the periphery of the body is filtered in the thalamus
before reaching the cortex.
The thalamus is divided in multiple specialised nuclei. The thalamic nuclei which process
somatosensory information are the ventro posterolateral nucleus (VPL), which processes information
from the body, and the ventro posteromedial nucleus (VPM), which processes information from the
head. Somatosensory information also reaches the intralaminar nuclei in the thalamus which are
inespecific: they process mixed type of information and they reach several different cortical areas.

The somatosensory cortex

In 1909 Korbinian Brodmann defined different areas of the cerebral cortex based on
their cytoarchitecture (structure and shape of the cells). He discovered that different areas of the
cortex have different types, density and organization of neurons and labelled each area with a
number. The majority of the areas that Brodmann defined have been correlated with specific
physiological functions. His terminology is still in current use today.
In the case of the somatosensory cortices, Brodmanns areas 1,2 and 3 correspond to the primary
somatosensory cortex (SI) and area 7 corresponds to the associative somatosensory cortex.
The somatosensory cortex is divided into:

Primary somatosensory cortex (SI) where somatosensory

information is still segreated. It is arranged by receptor type and
location of origin.

Secondary somatosensory cortex (SII) where somatosensory

information is integrated.

Somatosensory associative areas, where information from

different sensory modalities is further integrated. The
somatosensory associative cortices are found in the parietal
cortex. The posterior parietal cortex is an area where several
segregated streams of sensory information converge to produce
a complex representation. The posterior parietal cortex
processes somatic sensation together with visual information
and a persons state of attentiveness.

All the somatosensory cortices are part of the neocortex. Like most of the neocortex, they too are
divided in six layers, layer I being at the surface and layer VI the deepest one. Each cortical layer is
characterised by different types of neurons and specific connectivity.
Thalamic input goes into layer IV and a minor input goes to layer VI. Information from layer IV is then
transmitted to layer II and III where some connections travel horizontally, or intracortically, to
neighbouring cortical areas. Apical dentrites from layer V cells contact layer III small pyramidal
neurons and send outputs from the somatosensory cortex to subcortical areas. Finally information
from layer VI that comes from layer V and the thalamus, sends projections back to the thalamus. This
forms a loop of inputs and outputs of information coming into the cortex from deeper nuclei and being
sent back to the thalamus and other neighbouring cortical areas.

The somatosensory cortices

Different types of neurons in the primary somatosensory cortex

SI cortical connections

A topographic map in SI
The term somatotopy refers to the correspondence between a receptors in the body and the
dedicated area of the cortex that is activated by it. In the case of the somatosensory system, this
means that for every part of the body there is a corresponding area in the brain. Atopographic
map of the whole body can therefore be found in the somatosensory cortex. The sensory topographic
map of our body is known as the sensory homunculus, and it represents the location and the
amount of cortical area dedicated to a particular part of the body or the head.
As you can see from the figure, the representation of some parts of the body and head are distorted in
the somatosensory cortex. Certain body parts, like the lips and the hands, have more cortical area
dedicated to them than some others. This is because spatial resolution in the cortex is correlated with
the innervation density of the skin, so the more sensitive a region of the body is, the more space it will
take in the cortex. You could look at the sensory homunculus as a map of how sensitive our bodies

are: hands and lips are clearly the most sensitive areas, whereas legs and torso are much less
sensitive.
The sensory homunculus is different in each species. Rabbits have a broad cortical area dedicated to
the representation of their face and especially their teeth. Cats have a great representation of their
heads and their four paws. Most remarkably rats and mice have a large cortical area completely
dedicated only to their whiskers, known as the barrel cortex. This is such a sophisticated example of
cortical specialisation that it has become one of the principal models for the investigation of the
somatosensory system.
Somatosensory maps are plastic and can be modified by training or use. They are shaped by
experience during development and can be modified during adulthood. As reported by Mezernich in
1984, if a monkey looses a finger of the hand, the receptive field of the monkey's brain adjusts, so the
somatosensory homunculus allocates more of its receptive field to the adjacent fingers.

The humunculus

Pathologies of the somatosensory system: the phantom limb

Early reported cases:
Ambroise Pare (1510-1590) was a barber who became a French military surgeon. He observed that
amputees reported strange sympthoms. In the 16th century he wrote: "...the patients who many
months after the cutting away of the leg grievously complained that they yet felt exceeding great pain
of the leg so cut off..."
Silas Weir Mitchell (1871) who was a surgeron in the American Civil War wrote: "...nearly every man
who loses a limb, carries about with him a constant or inconstant phantom of the missing member, a
sensory ghost of that much of himself and sometimes a most inconvenient presence faintly felt at
times, but ready to be called up to his perceptios by a blow, touch or a wind of change...".
What we know today about the phantom limb
Phantom limbs occur in nearly every case of amputees who lose a limb.
The phantom is usually described as resembling the normal somatosensory experience of having the
physical limb, the same way as it was before being amputated. It might move, the same way as

before, with normal shape and size. As time passes after the amputation, the shape and size of the
phantom can become distorted. This is termed as the telescopic phenomenon of the phantom limb.
Phantom limbs can sometimes be painful and the sensation can range from itch to a strong muscular
contraction to burning pain. Not only this, but pain to the missing limb can be elicited by stimulation of
a different part of the body.
Why do amputees feel sensations on a limb that doesn't exist?
When a limb is amputated, changes occur both in the peripheral and the central nervous system:
The fibres belonging to the somatic receptors that innervated the amputated limb are cut, turning into
free nerve endings that will eventually loose the myelination. This doesn't mean that they stop
transmitting pulses, quite the opposite, the free nerve endings are activated and fire every time the
stump is stimulated.
However, the brain codes information coming from those fibers as if they were still coming from the
missing limb. So the brain constructs a percept with that sensation that corresponds to the phantom.
In the somatosensory cortex, the areas dedicated to the missing limb do not receive as much
stimulation anymore. When an area in the brain stops receiving information it eventually degenerates.
As explained earlier, adjacent areas in the homunculus take over the unused area to make the most of
the cortical space. It can therefore happen that stimulation of one body part can provoke sensations
on the phantom limb.
If you would like to find out more about the phantom limb, we recommend the book by V.S.
Ramanchandran and S Blakeslee: "Phantoms in the brain".

Summary

The somatosensory system processes mechanoreceptive,

proprioceptive, nociceptive and information related to
temperature.

The somatosensory system carries information from the

periphery to the cerebral cortex through the lemniscal,
anterolateral and trigeminal pathways. All sensory inputs cross
to the midline, so the information coming from the left part of the
body is processed by the right hemisphere of the brain and
viceversa.

The somatosensory system maintains its somatotopy along its

pathways from the periphery to the cerebral cortex.

The somatosensory cortex is part of the neocortex. It is divided

in six layers, each of which have different types of neurons
forming different connections.

There is a sensory homunculus in the somatosensory cortex

that represents our body and head. The amount of cortical area
dedicated to each part will determine how sensitive that area is.

Further readings
References

Phillips, J. R., Johansson, R.S., Johnson, K.O. 1990.

Representation of braille characters in human nerve fibres.
Experimental brain research, 81:589-592.

Mountcastle, V. 1957. Modality and topographic properties of

single neurons of cat's somatic sensory cortex. J
Neurosci,20:408-434.

Lbke, J., Feldmeyer, D. 2007. Excitatory signal flow and

connectivity in a cortical column: focus on the barrel
cortex. Brain Struct Funct 212(1):3-17.

Buonomano, D.V., Merzenich, M.M. 1998. Cortical plasticity:

from synapses to maps. Annual reviews neuroscience. 21:149186.

Merzenich, M.M., Nelson, R.J., Stryker, M.P., Cynader, M.S.,

Schoppmann, A., Zook, J.M. 1984. Somatosensory cortical map
changes following digit amputation in adult monkeys. J Comp
Neurol. 224(4):591-605.

Woodhouse, A. 2005. The Phantom limb sensation. Clin Exp

Pharmacol Physiol. 32(1-2):132-4

Flor, H., Nikolajsen, L., Jensen, T.S.. 2006. The Phantom limb
pain: a case of maladaptive CNS plasticity?. Nat Neurosci Rev.
7(11):873-81.

Chapter 12: The Somatic Sensory System In: Neuroscience:

Exploring the Brain (Bear MF, Connors BW, Paradiso BW., eds).
Williams & Wilkins.

Chapter 18: The Functional Organization of Perception and

Movement; Chapter 22: The Body Senses In: Principles of
Neuroscience (Kandel ER, Schwartz JH and Jessel TM., eds).
McGraw-Hill.

Chapter 23: Touch. In: Principles of Neuroscience (Kandel ER,

Schwartz JH and Jessel TM., eds). McGraw-Hill.

Bibliography

Chapter 26: The Somatic Sensation. In: The Fundamental

Neuroscience Ed. (Zigmond MJ, Bloom FE, Landis SC, Roberts
JL and Squire LR., eds). Academic Press.

CELLULAR & MOLECULAR

Action potentials and synaptic transmission

Ion channels and membrane potential
Neurotransmitters and their receptors
Overview of the neuron
NS ANATOMY

Central nervous system

Peripheral nervous system
DEVELOPMENT OF NS

Axonal growth
Cell differentiation
Induction and pattern formation
Neural Crest
Programmed cell death
Synapse formation
NEUROPHARMACOLOGY

Anaesthetics and Analgesics

Antidepressants
Antipsychotics
Anxiolytics
Drug abuse
Neurobiology of Pain

Stimulants
LEARNING AND MEMORY

Declarative memory
Emotional memory
Explicit and implicit memory
Memory consolidation
Molecular basis of memory
Overview of the hippocampus
Procedural memory
Spatial learning
NEUROENDOCRINOLOGY

Oxytocin and vasopressin

Sexual behaviour and competition
Sexual development & dimorphism
Sleep and hunger
Stress
The brain and it's hormones
SENSORY SYSTEMS

Audition
Gustation
Olfaction
Somatosensation
Vision
NEUROIMMUNOLOGY

Brain repair after acute inflammation

Clinical assessment of neuroinflammation

Early response to infection and injury
Immune cells and mediators in the brain
Neural control of host defense
System inflammation and the development of neurode
The role of the immune system in the healthy brain
COMPUTATIONAL NS

Artificial neural nets

Neural noise
Noisy but reliable computation
Population codes
Single neuron codes
Spike triggered average
The neural code
DISEASES OF THE CNS

Alzheimer's disease
Dementia
Depression
Epilepsy
Motor neuron disease
Multiple sclerosis
Parkinson's disease
Schizophrenia
Stroke
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Somatosensory Receptors
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There are various types of tactile

mechanoreceptors that work together to signal and
process "touch."
LEARNING OBJECTIVE[ EDIT ]

Describe the structure and function of mechanoreceptors

KEY POINTS[ EDIT ]

The four major types of tactile mechanoreceptors include: Merkel's disks,

Meissner's corpuscles, Ruffini endings, and Pacinian corpuscles.

Merkel's disk are slow-adapting, unencapsulated nerve endings that respond

to light touch; they are present in the upper layers of skin that has hair or
is glabrous.

Meissner's corpuscles are rapidly-adapting, encapsulated neurons that

responds to low-frequency vibrations and fine touch; they are located in the
glabrous skin on fingertips and eyelids.

Ruffini endings are slow adapting, encapsulated receptors that respond to

skin stretch and are present in both the glabrous and hairy skin.

-Pacinian corpuscles are rapidly-adapting, deep receptors that respond to

deep pressure and high-frequency vibration.

TERMS[ EDIT ]

glabrous
smooth, hairless, bald

dendrite
branched projections of a neuron that conduct the impulses received from other
neural cells to the cell body
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FULL TEXT[ EDIT ]

Somatosensory Receptors
Sensory receptors are classified into five
categories:mechanoreceptors, thermoreceptors, proprioceptors, pain
receptors, and chemoreceptors. These categories are based on the nature of
the stimuli that each receptor class transduces. Mechanoreceptors in the
skin are described as encapsulated or unencapsulated. A free nerve ending
is an unencapsulated dendrite of a sensory neuron; they are the most
common nerve endings in skin. Free nerve endings are sensitive to painful
stimuli, to hot and cold, and to light touch. They are slow to adjust to a
stimulus and so are less sensitive to abrupt changes in stimulation.

Mechanoreceptors

There are three classes of mechanoreceptors: tactile, proprioceptors, and

baroreceptors. Mechanoreceptors sense stimuli due to physical
deformation of theirplasma membranes. They contain mechanicallygated ion channels whose gates open or close in response to pressure,
touch, stretching, and sound. There are four primary tactile
mechanoreceptors in human skin: Merkel's disks, Meissner's corpuscles,
Ruffini endings, and Pacinian corpuscle; two are located toward the surface
of the skin and two are located deeper . A fifth type of mechanoreceptor,
Krause end bulbs, are found only in specialized regions.

Primary mechanoreceptors
Four of the primary mechanoreceptors in human skin are shown. Merkel's disks, which
are unencapsulated, respond to light touch. Meissner's corpuscles, Ruffini endings,
Pacinian corpuscles, and Krause end bulbs are all encapsulated. Meissner's corpuscles
respond to touch and low-frequency vibration. Ruffini endings detect stretch, deformation
within joints, and warmth. Pacinian corpuscles detect transient pressure and highfrequency vibration. Krause end bulbs detect cold.

Merkel's disks are found in the upper layers of skin near the base of the
epidermis, both in skin that has hair and on glabrous skin; that is, the
hairless skin found on the palms and fingers, the soles of the feet, and the

lips of humans and other primates. Merkel's disks are densely distributed
in the fingertips and lips. They are slow-adapting, unencapsulated nerve
endings, which respond to light touch. Light touch, also known as
discriminative touch, is a light pressure that allows the location of a
stimulus to be pinpointed. The receptive fields of Merkel's disks are small,
with well-defined borders. That makes them very sensitive to edges; they
come into use in tasks such as typing on a keyboard.
Meissner's corpuscles, also known as tactile corpuscles, are found in the
upper dermis, but they project into the epidermis . They are found
primarily in the glabrous skin on the fingertips and eyelids. They respond
to fine touch and pressure, but they also respond to low-frequency
vibration or flutter. They are rapidly- adapting, fluid-filled, encapsulated
neurons with small, well-defined borders which are responsive to fine
details. Merkel's disks and Meissner's corpuscles are not as plentiful in the
palms as they are in the fingertips.

Meissner corpuscles
Meissner corpuscles in the fingertips, such as the one viewed here using bright field light
microscopy, allow for touch discrimination of fine detail.

Deeper in the epidermis, near the base, are Ruffini endings, which are also
known as bulbous corpuscles. They are found in both glabrous and hairy
skin. These are slow-adapting, encapsulated mechanoreceptors that detect
skin stretch and deformations within joints; they provide valuable feedback
for gripping objects and controlling finger position and movement. Thus,
they also contribute to proprioception and kinesthesia. Ruffini endings
also detect warmth. Note that these warmth detectors are situated deeper in
the skin than are the cold detectors. It is not surprising, then, that humans
detect cold stimuli before they detect warm stimuli.

Pacinian corpuscles, located deep in the dermis of both glabrous and hairy
skin, are structurally similar to Meissner's corpuscles. They are found in the
bone periosteum, joint capsules, pancreas and other viscera, breast, and
genitals . They are rapidly-adapting mechanoreceptors that sense deep,
transient (not prolonged) pressure, and high-frequency vibration. Pacinian
receptors detect pressure and vibration by being compressed which
stimulates their internal dendrites. There are fewer Pacinian corpuscles and
Ruffini endings in skin than there are Merkel's disks and Meissner's
corpuscles.

Pacinian corpuscles

Pacinian corpuscles, such as these visualized using bright field light microscopy, detect
pressure (touch) and high-frequency vibration.
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Boundless Biology
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Which type of mechanoreceptor is CORRECTLY paired with its description?

Meissner's corpuscles: slow-adapting, encapsulated neurons that respond to
discriminative touch, Ruffini endings: fast-adapting, encapsulated neurons that respond
to skin deformations, Pacinian corpuscles: slow-adapting, deeply-located neurons that
respond to deep pressure, and Merkel's disks: slow-adapting, unencapsulated nerve
endings that reponse to discriminative touch

KEY TERM REFERENCE

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Source: Boundless. Somatosensory Receptors. Boundless Biology. Boundless, 24

Mar. 2015. Retrieved 20 May. 2015
from https://www.boundless.com/biology/textbooks/boundless-biologytextbook/sensory-systems-36/somatosensation-206/somatosensory-receptors-77812012/

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Source: Boundless. Somatosensory Receptors. Boundless Biology. Boundless, 24 Mar. 2015.

Retrieved 26 May. 2015 from https://www.boundless.com/biology/textbooks/boundless-biologytextbook/sensory-systems-36/somatosensation-206/somatosensory-receptors-778-12012/