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Table I
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
Sub-Sub Modality
touch
pressure
flutter
vibration
muscle length
muscle tension
joint pressure
muscle length
muscle tension
joint pressure
joint angle
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.
tests whether the proprioceptive components are working properly when the visual
cues are missing and proprioceptive cues are the major sources of information.
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.
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.
Figure 2.5
The
locations of
somatosens
ory
receptors in
the body.
Figure 2.6
The
locations of
cutaneous
(somatosens
ory)
receptors in
hairy and
non-hairy
(glabrous)
skin.
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
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.
Figure 2.11
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.
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.
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
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.
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.
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
Signals
Pacinian
corpuscle
Ruffini
corpuscle
Encapsulated
& layered
Encapsulated
collagen
Hair follicle
Merkel
complex
Free Nerve
Ending
Touch: Vibration
Frequency: 100-300 Hz
Unencapsulated
Touch: Movement
Direction &
Velocity
Specialized
epithelial cell
Unencapsulated
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,
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.
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:
* 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.
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.
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:
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.
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 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.
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:
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.
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
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
Further readings
References
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.
Bibliography
Axonal growth
Cell differentiation
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Somatosensation
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Somatosensory Receptors
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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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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
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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