Lect. univ. dr.

Loredana - Cristina MEREU

Laboratory of Biophysics & Med. Physics, Faculty of Physics,
'Alexandru Ioan Cuza' University of Iasi

I.

General principles of sensory physiology

II.

The somatosensory System

III.
IV.
V.

Chemical Senses: Taste and Olfaction

Vision
Hearing and Equilibrium

SOMATIC SENSES
The somatic sensory system has two major
components:
I. General somatic include touch, pain, vibration,
pressure, temperature
II. Proprioceptive detect stretch in tendons and
muscle provide information on body position,
orientation and movement of body in space.
Together, these two subsystems give humans
and other animals the ability to identify the shapes and
textures of objects, to monitor the internal and external
forces acting on the body at any moment, and to detect
potentially harmful circumstances.

The somatosensory system is subdivided into:

1. Exteroreceptors - these are the familiar receptors
found in the skin that mediate the sub-modalities of
touch, pain and temperature.
These types of sensory input can mediate both
rapid responses (e.g. reflexes) and reach the cerebral
cortex and induce perception.
2. Interoreceptors - are found in internal organs and
convey signals that include distension of the stomach,
carbon dioxide concentration in the blood etc. This type
of sensory input is obviously of great importance but
difficult to study experimentally (in the CNS).

The somatosensory system is subdivided into:

3. Proprioceptors.
Proprioceptor afferents are found in muscles and
at joints; they mediate the detection of muscle stretch
and the degree of extension at a joint.
For the most part, this type of sensory input is
involved in motor control rather than perception.
The two major classes of proprioceptive input
come from:
(a) Golgi tendon organs
(b) muscle spindles.

All Somatic Sensation Begins with Receptors and

Ganglion Cells
The components of the
somatic
sensory
system
process information conveyed
by mechanical stimuli that
impinge upon the body surface
or that are generated within the
body itself (proprioception).
This
processing
is
performed
by
neurons
distributed across several brain
structures that are connected
by
both
ascending
and
descending pathways.

All Somatic Sensation Begins with Receptors and

Ganglion Cells
This activity is conveyed centrally via a chain of
neurons, referred to as the first-, second-, and thirdorder cells.
First-order neurons are
located in the dorsal root
and cranial nerve ganglia.
Second-order neurons
are located in brainstem
nuclei.
Third-order neurons are
found in the thalamus,
from where they project
to the cerebral cortex.

All Somatic Sensation Begins with Receptors and

Ganglion Cells
Transmission of afferent mechanosensory
information from the periphery to the brain begins with
a variety of receptor types that initiate action
potentials.

These pathways are topographically arranged

throughout the system, the amount of cortical and
subcortical space allocated to various body parts being
proportional to the density of peripheral receptors.

Ascending Pathways
The afferent fibers from the receptors for each of
these sensations are projected to the dorsal horn of the
spinal cord where they synapse with the dorsal root
ganglion. From there the fibers split into two pathways:
(a) The medial
lemniscal
tract
travels in the dorsal
column and carries
tactile
sensation
and proprioceptive
information to the
thalamus and then
to
the
somatosensory
cortex.

Ascending Pathways
(b)The spinothalamic tract travels contralaterally up the
spinal cord in the anterolateral column and carries painful
and thermal sensations to the thalamus and finally to the
somatosensory cortex.

Somatotopic map

The sensory information

is
kept
somatotopicaly
organized throughout the entire
pathway for each of the tracts.

(A) The human somatotopic map first defined in the

1930s has remained generally valid.
(B) Diagram along the
plane in (A) showing the
somatotopic
representation of body
parts from medial to
lateral.
(C) Homunculus - the
amount
of
somatic
sensory cortex devoted
to the hands and face is
much larger than the
relative amount of body
surface in these regions.

The innervation arising from a single dorsal root

ganglion and its spinal nerve is called a dermatome.
In humans, the
cutaneous area of each
dermatome has been
defined in patients in
whom specific dorsal
roots were affected or
after
surgical
interruption (for relief of
pain or other reasons).
Such studies show that
dermatomal maps vary
among individuals.

Dermatomal maps
Despite these limitations, knowledge of
dermatomes is essential in the clinical evaluation of
neurological patients, particularly in determining the
level of a spinal lesion.

Sagittal view of the spinal cord showing the origin of nerves

How it works
Despite their variety, all somatic sensory receptors
work in fundamentally the same way:
Stimuli applied to the skin deform or otherwise
change the nerve endings, which in turn affects the
ionic permeability of the receptor cell membrane.
Changes in permeability generate a depolarizing
current in the nerve ending, thus producing a receptor
(or generator) potential that triggers action potentials.
This overall process, in which the energy of a
stimulus is converted into an electrical signal in the
sensory neuron, is called sensory transduction and is
the critical first step in all sensory processing.

How it works
The quality of a mechanosensory (or any other)
stimulus (i.e., what it represents and where it is) is
determined by the properties of the relevant receptors
and the location of their central targets.
The quantity or strength of the stimulus is
conveyed by the rate of action potential discharge
triggered by the receptor potential (although this
relationship is nonlinear and often quite complex).
Some receptors fire rapidly when a stimulus is first
presented and then fall silent in the presence of
continued stimulation (which is to say they adapt to
the stimulus), whereas others generate a sustained
discharge in the presence of an ongoing stimulus.

How it works
The usefulness of having some receptors that
adapt quickly and others that do not is to provide
information about both the dynamic and static
qualities of a stimulus.
Receptors that initially fire in the presence of a
stimulus and then become quiescent are particularly
effective in conveying information about changes in the
information the receptor reports; conversely, receptors
that continue to fire convey information about the
persistence of a stimulus.

How it works
Accordingly, somatic sensory receptors and the
neurons that give rise to them are usually classified into
rapidly or slowly adapting types.
Rapidly adapting, or phasic
receptors respond maximally
but briefly to stimuli; their
response decreases if the
stimulus is maintained.
Conversely, slowly adapting,
or tonic receptors keep firing
as long as the stimulus is
present.

How it works
An applied pressure on the skin (e.g. pushing on a
key on a computer keyboard, or sitting on a hard
surface) distorts the concentric layers. This results in
deformation of the cell membrane of the sensory
neurone. Deformation opens the ion channels in the
membrane (changes the shape of the protein slightly) so
Na+ ions pass into the neurone.

How it works
This produces a depolarisation called a generator
potential of about 1mV across the membrane. If the
generator potential exceeds the threshold it triggers the
generation of action potentials at the first node of
Ranvier, nerve impulses are then transmitted along the
length of the sensory neurone to the CNS.

How it works
When the pressure stimulus is removed the
corpuscle resumes its normal shape producing another
transitory deformation of the receptor membrane in the
process and another brief generator potential, which will
also generate action potentials to indicate the pressure,
has been removed.
The size of the generator potential is
proportional to the amount of opening of the ion
channels, which is proportional to the amount of
deformation, which is proportional to the intensity of
the stimulus, so the size of the generator potential is
proportional to the intensity of the stimulus (called a
graded response).

How it works
The number of action potentials generated is
proportional to the size of the generator potential, the
stronger the stimulus the bigger the generator potential
the more action potentials are generated (a frequency
coded response).

Sensory Information Is Transmitted Along

Labeled Lines
In one particular population of somatosensory
neurons, activity is always interpreted by the CNS as a
painful stimulus, no matter whether the stimulus is
natural (a sharp instrument) or artificial (electrical
stimulation of the appropriate axons). An entirely
separate population of neurons (colored blue) would
signal light pressure.
Why this is so can be seen
from the fact that receptors
are selective not only in what
drives them, but also in the
postsynaptic
targets
with
which they communicate.

Sensory Information Is Transmitted Along

Labeled Lines
Each ganglion cell transmits its activity into a welldefined region of the CNS, after which a strictly
organized series of synaptic connections relays
information in a sequence that eventually leads to the
thalamus and then to the cerebral cortex.
It is this orderly relay
from receptor to ganglion cell
to central neurons at each of
several stations that makes up
a labeled line.

Sensory Information Is Transmitted Along

Labeled Lines
All sensory information arising from a single class
of receptors is referred to as a modality (e.g., the
sensations of pain and light pressure involve distinct
modalities).
Thus,
the
existence of labeled
lines means that
neurons in sensory
systems
carry
specific modalities.

Somatic Receptors are divided into two groups:

I. Free or Un-encapsulated nerve endings
- are abundant in epithelia and underlying connective
tissue (nociceptors
and thermoreceptors); Two
specialized types of free nerve endings:

Merkel discs
lie
in
the
epidermis, slowly
adapting
receptors for light
touch
Hair
follicle
receptors

Rapidly adapting
receptors
that
wrap around hair
follicles

Somatic Receptors are divided into two groups:

II. Encapsulated nerve endings - consist of one or
more neural end fibers enclosed in connective tissue:

Meissners corpuscles; Pacinian corpuscles;

corpuscles
Proprioceptors:
Muscle spindles monitors the
changing length of a muscle,
imbedded
between
muscle
fascicles
Golgi tendon organs located
near the muscle-tendon junction,
monitor tension within tendons
Joint
kinesthetic receptors sensory nerve endings within the
joint capsules, sense pressure and
position.

Ruffinis

In the 1920s and 1930s, there was a virtual

industry classifying axons according to their
conduction velocity. Three main categories were
discerned, called A, B, and C.
A comprises the
largest and fastest
axons,
C the smallest
and slowest.
Mechanoreceptor
axons generally fall
into category A.

Classifying axons according to their conduction velocity.

The A group is further broken down into three

subgroups designated:
alpha (the fastest)
betha
delta (the slowest)
To make matters even more
confusing, muscle afferent
axons are usually classified
into
four
additional
groups: I (the fastest),
II,
III,
and IV (the slowest)with
subgroups designated by
lowercase roman letters!

Somatic Receptors

Somatic Receptors
The receptors that transduce light or sound are
highly specialized cells; in contrast, exteroreceptors
are merely nerve endings in the skin.
There are many forms of exteroreceptors.
Based on function, this variety of somatic receptors
can be divided into three groups:
1.

Mechanoreceptors respond to mechanic stimuli

2.

Nociceptors - respond to pain

3.

Thermoceptors - respond to temperature

Somatic Receptors
The most important distinction is between:
a. Touch or mechano-receptors - the nerve
endings are usually associated with some specialized
structures such as hairs. The axons of touch fibers are
heavily myelinated allowing them to conduct action
potentials rapidly. The response of touch fibers
depends on their associated structures.
b. Pain and temperature receptors. The nerve
endings of pain and temperature receptors are simple
and the axons are either unmyelinated (C type) or
lightly myelinated.

The Skin Harbors Morphologically Distinct

Mechanoreceptors
Mechanoreceptors detect mechanical energy,
typically consist of ion channels linked to external cell
structures (i.e. hairs) & internal structures (i.e.
cytoskeleton). Bending/stretching plasma membrane
changes permeability to sodium & potassium ions.

Simulated activity patterns in different

mechanosensory afferents as Braille is read.

The Skin Harbors Morphologically Distinct

Mechanoreceptors
Four major types of mechanoreceptors are
specialized to provide information to the central nervous
system about touch, pressure, vibration, and
cutaneous tension:
Meissners corpuscles;
Pacinian corpuscles;
Merkels disks;
Ruffinis corpuscles.

Mechanoreceptors
These receptors are referred to collectively as
low-threshold
(or
high-sensitivity)
mechanoreceptors because even weak mechanical stimulation
of the skin induces them to produce action potentials
and they are innervated by relatively large myelinated
axons (type A), ensuring the rapid central transmission
of tactile information.

Touch Receptors
Free nerve endings Simplest mechanoreceptors
are in skin; touch pressure & pain (nociceptors)
More specialized tactile receptors
Merkel Discs- touch and pressure
Meissner corpuscles- Light pressure
Ruffini corpuscles- Heavy pressure
Pacinian corpuscles- subcutaneous- vibration
Other Mechanoreceptors
Proprioreceptors
Muscle spindles- Stretch receptors
Golgi tendon organs- Force generated by
muscle
Blood Pressure Measured at carotid sinus and
aortic arch; Baroreceptors- Tension in walls of
vessels

Tactile Discrimination
Sensitivity of Tactile Discrimination Varies with
Location on the Body Surface
There are differences in mechanosensory
discrimination across the body surface.
The accuracy of our sense of touch is not the same
all over the body.
Fingers can distinguish things 2 mm apart, forearms
40 mm apart.
Mechanosensory receptors are more numerous in
finger tips and have smaller receptive fields.

Tactile Discrimination

Sensitivity of
Tactile
Discriminati
on
Varies
with Location
on the Body
Surface

Cutaneous Receptors
1. Touch Receptors: fine touch
a. Meissners corpuscle fine touch,
discrimination; found concentrated in places where you
need to have a lot of responsiveness to a little input.
b. Merkel disks - found deep at the junction of the
epidermis and dermis.
c. Root hair plexus - at the base of hair follicles.

Cutaneous Receptors
2. Touch Receptors: pressure sensitive
a. Ruffinis endings and Krause's end bulbs
encapsulated
pressure
sensors,
dermis
(and
elsewhere), respond to continuous pressure
b. Pacinian corpuscles deep pressure sensors,
onion shaped capsule (layers of Schwann cells enclosed
in a connective tissue membrane), respond to on-off
pressure or vibration.
3. Temperature Free nerve endings, some
responsive to heat and others responsive to cold
4. Pain - Free nerve endings, respond to chemicals
released from damaged tissues.

Encapsulated Nerve Endings-Meissners corpuscles

Lie between the dermal papillae just beneath the
epidermis of the fingers, palms, and soles. Meissners
corpuscles are the most common mechanoreceptors of
glabrous (smooth, hairless) skin (the fingertips, for
instance), and their afferent fibers account for about
40% of the sensory innervation of the human hand.
These corpuscles are
particularly efficient in
transducing information
about the relatively lowfrequency vibrations (30
50 Hz) that occur when
textured
objects
are
moved across the skin.

Encapsulated Nerve Endings Meissners corpuscles

How it works
Are elongated receptors formed by a connective
tissue capsule that comprises several lamellae of
Schwann cells. The center of the capsule contains one
or more afferent nerve fibers that generate rapidly
adapting action potentials following minimal skin
depression.
When a force is
applied to the dermal
papilla
containing
the
Meissner corpuscle, the
laminar
cells
in
the
corpuscle slide past one
another.

Encapsulated Nerve Endings Meissners corpuscles

How it works
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.

Example of a simple receptor - Pacinian

corpuscles - are large encapsulated endings located in
the subcutaneous tissue (and more deeply in
interosseous membranes and mesenteries of the gut).

They make
up 1015%
of
the
cutaneous
receptors in
the hand.
These receptors differ from Meissners corpuscles in
their morphology, distribution, and response threshold.

How it works
The Pacinian corpuscle has an onion-like
capsule in which the inner core of membrane lamellae is
separated from an outer lamella by a fluid-filled space.
One or more rapidly adapting afferent axons lie at the
center of this structure.

The capsule acts as a filter, in this case allowing

only transient disturbances at high frequencies (250
350 Hz) to activate the nerve endings.

How it works

Pacinian corpuscles adapt more rapidly than Meissners

corpuscles and have a lower response threshold. These
attributes suggest that Pacinian corpuscles are involved in
the discrimination of fine surface textures or other moving
stimuli that produce high-frequency vibration of the skin.
In corroboration of this supposition, stimulation of Pacinian
corpuscle afferent fibers in humans induces a sensation of
vibration or tickle.
Because they are
rapidly adapting, Pacinian
corpuscles,
like
Meissners corpuscles,
provide
information
primarily
about
the
dynamic qualities of
mechanical stimuli.

Slowly adapting cutaneous mechanoreceptors

include Merkels disks and Ruffinis corpuscles

Merkels disks are located in the epidermis,

where they are precisely aligned with the papillae that lie
beneath the dermal ridges.
They account for
about 25% of the
mechanoreceptors
of the hand and are
particularly dense in
the fingertips, lips,
and
external
genitalia.

How it works
The slowly adapting nerve fiber
associated with each Merkels disk
enlarges into a saucer-shaped ending
that is closely applied to another
specialized cell containing vesicles
that apparently release peptides that
modulate the nerve terminal. Selective
stimulation of these receptors in
humans produces a sensation of light
pressure. These several properties
have led to the supposition that
Merkels disks play a major role in the
static discrimination of shapes,
edges, and rough textures.

How it works
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, 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.

Ruffinis corpuscles, although structurally similar to

other tactile receptors, are not well understood. These
elongated, spindle-shaped capsular specializations are
located deep in the skin, as well as in ligaments and
tendons. The long axis of the corpuscle is usually
oriented parallel to the stretch lines in skin.
Thus, Ruffinis
corpuscles
are
particularly sensitive
to the cutaneous
stretching produced
by digit or limb
movements.
Ruffinis corpuscles account for about 20% of the
receptors in the human hand and do not elicit any particular
tactile sensation when stimulated electrically.

How it works

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.

The hair follicle receptor

is an
unencapsulated
cutaneous
receptor.

The 1 afferent terminal axons spiral around the hair follicle base
or run parallel to the hair shaft forming a lattice-like pattern. 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. 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. 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.

Response Properties of Fine Touch Fibers

The structure of the accessory tissue of the nerve
ending determines its response.
Meissners corpuscles fast adapting (FA) receptors
for discriminative touch
Pacinian corpuscles sensitive to deep pressure
fast (FA) adapting receptors
are good for detecting
texture.
Ruffinis corpuscles Monitor continuous pressure
on the skin adapt slowly
(SA)- estimate the duration
of contact.

Response Properties of Fine Touch Fibers

Both the FA and SA afferent classes can be

subdivided on the basis of other aspects of their
receptive fields - defined as the region of skin from
which stimuli can evoke a response (i.e., change the
firing of the afferent axon).
Type 1 units have small
receptive fields with well-defined
borders. Particularly for glabrous
skin (i.e., hairless skin, such as
on the palms of the hands and
soles of the feet), the receptive
field has a circular or ovoid
shape, within which there is
relatively uniform and high
sensitivity
to
stimuli
that
decreases sharply at the border.

Receptive field characteristics for type 1 and

type 2 sensory afferents. Plots in the top row
show the threshold level of force needed to
evoke a response as a function of the distance
across the receptive field (shown on the hand)

Response Properties of Fine Touch Fibers

Type 1 units, particularly SA1 units, respond best to edges. That

is, a larger response is elicited from them when the edge of a
stimulus cuts through their receptive field than when the entire
receptive field is indented by the stimulus.
Type 2 units have wider
receptive fields with poorly
defined borders and only a single
point of maximal sensitivity, from
which there is a gradual reduction
in sensitivity with distance.
For comparison, a type 1
unit's receptive field typically will
cover approximately four papillary
ridges in the fingertip, whereas a
type 2 unit will have a receptive
field that covers most or all of a
finger.

Receptive field characteristics for type 1 and type

2 sensory afferents.

Response Properties of Fine Touch Fibers

The sensory systems encode four elementary

attributes of stimuli:
modality,
location,
intensity

and

timing,
which are manifested in
sensation.

The four attributes of sensation are illustrated in

this figure for the somatosensory modality of touch.

Response Properties of Fine Touch Fibers

A. In the human hand the submodalities of touch are

sensed by four types of mechanoreceptors. Specific
tactile sensations occur when distinct types of receptors
are activated. Firing of all four receptors produces the
sensation of contact with an object. Selective activation
of Merkel cells and Ruffini endings produces
sensations of steady pressure on the skin above the
receptor. When the same patterns of firing occur only in
Meissner's and Pacinian corpuscles, the tingling
sensation of vibration is perceived.

Response Properties of Fine Touch Fibers

B. Location and other spatial properties of a stimulus

are encoded by the spatial distribution of the population
of activated receptors. Each receptor fires action
potentials only when the skin close to its sensory
terminals is touched, i.e., when a stimulus impinges on
the receptor's receptive field. The receptive fields of
mechanoreceptors - shown as red areas on the finger tip
- differ in size and response to touch. Merkel cells and
Meissner's corpuscles provide the most precise
localization of touch, as they have the smallest receptive
fields and are also more sensitive to pressure applied by
a small probe.

Response Properties of Fine Touch Fibers

C. The intensity of stimulation is signaled by the firing

rates of individual receptors, and the duration of
stimulation is signaled by the time course of firing.
The spike trains below each finger indicate the
action potentials evoked by pressure from a small probe
at the center of the receptive field.
Two of these receptors
(Meissner's
and
Pacinian
corpuscles)
adapt
rapidly
to
constant
stimulation,
while the other two
adapt slowly.

Response Properties of Fine Touch Fibers

Mechanoreceptors in glabrous skin vary in the size and

structure of their receptive fields. Each colored area on
the hands indicates the receptive field of a different
sensory nerve fiber in the human median nerve.
A. The Merkel disk
receptor in the superficial
skin and the subcutaneous
Ruffini ending are slowly
adapting receptors. The
Merkel disk receptor has a
small,
highly
localized
receptive field, whereas
the Ruffini ending has a
large field (light purple)
with a central zone of
maximal sensitivity (dark
purple).

Depending
on
their
location,
individual Ruffini endings are excited
by stretch of the skin in specific
directions as indicated by arrows.

Response Properties of Fine Touch Fibers

B. The Meissner's corpuscle in the superficial skin and the subcutaneous

Pacinian corpuscle are rapidly adapting receptors. Meissner's corpuscles on
the fingertips have receptive fields averaging 2-3 mm in diameter, while
receptive fields on the palm average 10 mm in diameter. The receptive fields of
Pacinian corpuscles cover larger continuous surfaces on the fingers or palm
(light pink) but have a central zone of maximal sensitivity located directly above
the receptor (red).
C. Expanded view of the receptive
fields of mechanoreceptors in the skin.
The relative sensitivity to pressure is
shown as a contour map in which the
most sensitive regions are indicated in
red and the least sensitive areas in pale
pink. Receptive fields in the superficial
layers of the skin have many points of
high sensitivity, marking the positions of
the Meissner's corpuscles or Merkel
disk receptors. Receptive fields in the
deep layers have a single point of
maximal sensitivity overlying the
Pacinian or Ruffini receptor.

Response Properties of Fine Touch Fibers

The distribution of receptor types in the human hand

varies.

The
number of
sensory
nerve
fibers
innervating
an area is indicated
by the stippling
density, with the
highest density of
receptors shown by
the
heaviest
stippling.

Meissner's corpuscles (RA) and Merkel disk receptors (SA I)

are the most numerous receptors; they are distributed
preferentially on the distal half of the fingertip. Pacinian
corpuscles (PC) and Ruffini endings (SA II) are much less
common; they are distributed more uniformly on the hand,
showing little differentiation of the distal and proximal regions.

Response Properties of Fine Touch Fibers

The fingertips are the most densely innervated region of

skin in the human body, receiving approximately 300
mechanoreceptive nerve fibers per square centimeter.

The number of mechanoreceptive fibers is reduced to

120/cm2 in the proximal phalanges, and to 50/cm2 in the palm.

Response Properties of Fine Touch Fibers

Response Properties of Fine Touch Fibers

The shape and size of objects touching the hand

are encoded by populations of Merkel disk receptors.
A. The area of contact on
the skin determines the
total number of stimulated
Merkel disk receptors. The
pink region on the fingertip
shows
the
spread
of
excitation when probes of
different
diameters
are
pressed upon the skin with
constant force. The intensity
of color is proportional to the
firing rates of the stimulated
receptors.

Response Properties of Fine Touch Fibers

1. A small-diameter, sharp probe activates a small

population of Merkel receptors. However, the active
receptors fire intensely because all of the force is
concentrated at the small probe tip.
2. An intermediate-size probe
excites more receptors but the peak
firing rate in the population is
reduced. The probe does not feel as
sharp as the small-diameter probe.
3. A gently rounded, largediameter probe stimulates a large
population of receptors spread
across the width of the finger. These
receptors fire at low rates because
the force is spread over a larger
area of skin.

Response Properties of Fine Touch Fibers

The firing rate of individual Merkel disk receptors

signals the probe diameter.
B. These recordings of action potentials fired by a Merkel disk
receptor illustrate the responses evoked when probes of
decreasing size are pressed on the center of the receptive field.
All of the probes evoke a strong
initial response as contact is
made with the skin. The firing
rate of the neuron during steady
pressure is proportional to the
curvature of each probe. The
weakest responses are evoked
by flat surfaces and gently
rounded
(large
diameter)
probes. The firing rate increases
as the probe diameter becomes
smaller.

http://www.nature.com/nrn/journal/v12/n3/fig_tab/nrn2993_F1.html

Other Somatic Sensations - Warmth and Cold - Are

Mediated by Thermal Receptors
Skin temperature is coded by warmth receptors
and cold receptors.
A. Static temperatures. Cold receptors and warmth
receptors differ in the range of steady-state
temperatures to which they respond and in their peak
temperature sensitivities.
Cold receptors respond to steady-state
temperatures of 5-40C. Warmth receptors are
tonically active at steady temperatures of 2945C.
Cold receptors fire at highest rates at a
skin temperature of 25C, while warmth receptors
are most active at 45C.
At the normal skin temperature of 34C,
cold receptors are more active than warmth
receptors.

Other Somatic Sensations - Warmth and Cold - Are

Mediated by Thermal Receptors
B. Dynamic temperatures. Both receptors are more
sensitive to changes in skin temperature than to
constant temperatures.
Cooling the skin below the resting
level evokes a sharp rise in the firing rate of
cold receptors and silences warmth
receptors.
If the cold temperature is
maintained, the firing rates of the cold
receptors adapt. When the skin temperature
is rewarmed to the resting level, cold
receptors are briefly silenced, whereas
warmth receptors fire a burst of impulses.
Warming the skin produces the
opposite firing patterns in warmth and cold
receptors.

Dynamic Aspects of Somatic Sensory Receptive

Fields
When humans explore objects with their hands,
multiple contacts between the skin and the object
surface generate extraordinarily complex patterns of
tactile stimuli.
As a consequence, the somatic sensory system
must process signals that change continuously in
time.
Nonetheless, we routinely discriminate the size,
texture and shape of objects with great accuracy.

Dynamic Aspects of Somatic Sensory Receptive

Fields
Psychophysical analysis of tactile performance
suggests that something more than the cutaneous
periphery is needed to explain variations in tactile
perception.
For instance, even though we spend most of the
day wearing clothes, we usually ignore the tactile
stimulation that they produce. Some aspect of the
mechanosensory system allows us to filter out this
information and pay attention to it only when
necessary.

Dynamic Aspects of Somatic Sensory Receptive

Fields
The fascinating phenomenon of phantom limb
sensations after amputation provides further evidence
that tactile perception is not fully explained by the
peripheral information that travels centrally.

Dynamic Aspects of Somatic Sensory Receptive

Fields
The central nervous system clearly plays an
active role in determining the perception of the
mechanical forces that act on us.
Phantoms might simply be a curiosity - or a
provocative clue about higher order somatic sensory
processing - were it not for the fact that a substantial
number of amputees also develop phantom pain.

Dynamic Aspects of Somatic Sensory Receptive

Fields
This common problem is usually described as a
tingling or burning sensation in the missing part.
Sometimes, however, the sensation becomes a
more serious pain that patients find increasingly
debilitating.

Dynamic Aspects of
Somatic Sensory
Receptive Fields

Phantom pain
is, in fact, one of the
more common causes
of
chronic
pain
syndromes and is
extraordinarily difficult
to treat.

Mechanoreceptors Specialized for Proprioception

Whereas cutaneous mechanoreceptors provide
information derived from external stimuli, another major
class of receptors provides information about
mechanical forces arising from the body itself, the
musculoskeletal system in particular.
These are called proprioceptors,
roughly meaning receptors for self.
The purpose of proprioceptors
is primarily to give detailed and
continuous information about the
position of the limbs and other body
parts in space.

Mechanoreceptors Specialized for Proprioception

Whereas muscle spindles are specialized to
signal changes in muscle length (the degree to which
they
are
being
stretched),
low-threshold
mechanoreceptors in tendons, called Golgi tendon
organs, inform the central nervous system about
changes in muscle tension.
Finally, joint receptors are
rapidly
adapting
mechanoreceptors in and
around joints which gather
dynamic information about
limb position and joint
movement.

Muscle Spindles
Found in all but a few striated (skeletal) muscles.
Respond by causing muscle contraction
Each spindle is 3 to
10 mm long. Muscle
spindles consist of around
3 to 12 tiny specialized
intrafusal muscle fibers
surrounded by a capsule
of connective tissue.

The intrafusal fibers are distributed among the

ordinary (extrafusal) fibers of skeletal muscle in a
parallel arrangement.

Sensory Innervation of the Muscle Spindle - Central

region of each of these fibers has few or no actin and myosin
filaments. Therefore, this central portion does not contract
when the ends do. Instead, it functions as a sensory
receptor.

Motor Innervation of Muscle Spindle -The end portions

that do contract are excited by small gamma motor nerve
fibers (gamma efferent fibers) that originate from small type
A gamma motor neurons in the anterior horns of the spinal
cord. The large alpha efferent fibers (type A alpha nerve
fibers) innervate the extrafusal skeletal muscle.

Sensory Innervation of the Muscle Spindle

Muscle spindle receptor can be
excited in two ways:
1)

2)

lengthening the whole muscle

stretches the midportion of the
spindle and, therefore, excites
the receptor.
contraction of the end portions
of the spindle's intrafusal
fibers stretches the midportion
of the spindle and therefore
excites the receptor.

Sensory Innervation of the Muscle Spindle

Two types of sensory endings are found in this

central receptor area of the muscle spindle:
Primary Ending - this nerve fiber is a type Ia fiber
averaging 17 micrometers in diameter it transmits sensory
signals to the spinal cord at a velocity of 70 to 120 m/sec, as
rapidly as any type of nerve fiber in the entire body.
Secondary
Ending
usually one but sometimes
two smaller sensory nerve
fibers- type II fibers with
an average diameter of 8
micrometers-innervate the
receptor region on one or
both sides of the primary
ending.

Sensing Blood Pressure

Blood pressure is monitored at two main sites in
the body: carotid sinus, an enlargement of the left and
right internal carotid arteries, which supply blood to the
brain and the aortic arch, the portion of the aorta very
close to its emergence from the heart.
The walls of the blood
vessels at both sites
contain
a
highly
branched network of
afferent
neurons
called baroreceptors,
which detect tension
in the walls.

Sensing Blood Pressure

When the blood pressure decreases, the
frequency of impulses produced by the baroreceptors
decreases.
The CNS responds to this reduced input by
stimulating the sympathetic division of the autonomic
nervous system, causing an increase in heart rate and
vasoconstriction.
Both effects help to raise the blood pressure, thus
maintaining homeostasis.
A rise in blood pressure, conversely, reduces
sympathetic activity and stimulates the parasympathetic
division, slowing the heart and lowering the blood
pressure.

Sensing Blood Pressure

Mechanical distortion of the
plasma
membrane
of
mechanoreceptors
produces nerve impulses
that serve to monitor muscle
length from skeletal muscle
spindles and to monitor
blood
pressure
from
baroreceptors
within
arteries.

Golgi Tendon Organ Helps Control Muscle Tension

These mechanoreceptors, are encapsulated

sensory receptors innervated by branches of group Ib
afferents and are distributed among the collagen fibers
that form the tendons.
About 10 to 15 muscle
fibers
are
usually
connected to each Golgi
tendon
organ
is
stimulated when this small
bundle of muscle fibers is
"tensed" by contracting or
stretching the muscle.

Golgi Tendon Organ Helps Control Muscle Tension

Thus, the major difference in excitation of the

Golgi tendon organ versus the muscle spindle is
that the spindle detects muscle length and changes in
muscle length, whereas the tendon organ detects
muscle tension as reflected by the tension in itself.

Golgi Tendon Organ Helps Control Muscle Tension

The tendon organ, like the primary receptor of the

muscle spindle, has both a dynamic response and a
static response:

dynamic response: reacting intensely when the

muscle tension suddenly increases

static response: settling down within a fraction of a

second to a lower level of steady-state firing that is
almost directly proportional to the muscle tension.

Thus, Golgi tendon organs provide the nervous

system with instantaneous information on the degree
of tension in each small segment of each muscle.

Golgi Tendon Organ Helps Control Muscle Tension

Impulses from the tendon organ into the CNS are

transmitted through large, rapidly conducting type Ib
nerve fibers that average 16 micrometers in diameter.
When the Golgi tendon organs of
a muscle tendon are stimulated by
increased tension in the connecting
muscle, signals are transmitted to the
spinal cord to cause reflex effects in the
respective muscle. This reflex is
entirely inhibitory.

Inhibitory Nature of the Tendon Reflex and Its

Importance
Thus, this reflex provides a negative feedback
mechanism that prevents the development of too much
tension on the muscle and a protective mechanism to
prevent tearing of the muscle or avulsion of the tendon
from its attachments to the bone.

Joint Receptors
Joint receptors are found within the connective
tissue, capsule and ligaments of joints. 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.

Joint Receptors
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.

NOCICEPTION, THERMORECEPTION, AND ITCH

PAIN nocireceptors
Most of the sensory and somatosensory modalities
are primarily informative, whereas pain is a protective
modality.
Although similar in some ways to the sensory
processing of ordinary mechanical stimulation, the
perception of pain, called nociception, depends on
specifically dedicated receptors and pathways.
The relatively unspecialized nerve cell endings
that initiate the sensation of pain are called nociceptors
(noci is derived from the Latin nocere, to hurt).
Nociceptive means sensitive to noxious stimuli.
Noxious stimuli are stimuli that elicit tissue damage and
activate nociceptors.

PAIN nocireceptors
Nociceptors are sensory receptors -free (bare)
nerve endings found in the skin, muscle, joints, bone
and viscera, that detect signals from damaged tissue or
the threat of damage and indirectly also respond to
chemicals released from the damaged tissue.
Like other cutaneous and subcutaneous receptors,
they transduce a variety of stimuli into receptor
potentials, which in turn trigger afferent action potentials.

PAIN nocireceptors

Moreover, nociceptors, like other somatic sensory

receptors, arise from cell bodies in dorsal root ganglia (or in
the trigeminal ganglion) that send one axonal process to the
periphery and the other into the spinal cord or brainstem.
Because peripheral nociceptive axons terminate in
unspecialized free endings, it is conventional to categorize
nociceptors according to the properties of the axons
associated with them.

Different nociceptors/free nerve endings, and the fibers carrying pain

sensation from the nociceptors to the spinal cord.

PAIN nocireceptors

The axons associated with nociceptors, conduct

relatively slowly, being only lightly myelinated or, more
commonly, unmyelinated.

Accordingly, axons conveying information about pain

fall into:
A group of myelinated axons, which conduct at
about 20 m/s,
C fiber group of unmyelinated axons, which
conduct at velocities generally less than 2 m/s.

aA

- the largest and fastest axons: alpha (the fastest), betha, and delta (the slowest).
C -the smallest and slowest.
Muscle afferent axons: group I (the fastest), II, III, and IV (the slowest) - with subgroups a, b.

PAIN nocireceptors

Nociceptors are not uniformly sensitive. They

fall into several categories, depending on their
responses to mechanical, thermal, and/or chemical
stimulation liberated by the damage, tumor, and/or
inflammation.
Skin nociceptors may be divided into four
categories based on function:
I.

mechanonociceptors,

II. thermal nociceptors,

III. chemical nociceptors,
IV. polymodal nociceptors.

PAIN nocireceptors

The first type is termed high threshold

mechanonociceptors or specific nociceptors.
These faster-conducting A nociceptors respond
only to intense mechanical stimulation such as
pinching, cutting or stretching.
They are also excited by sharp objects that
penetrate, squeeze, or pinch the skin and therefore
mediate sensations of sharp or pricking pain.
Their
firing
rates
increase
with
the
destructiveness of mechanical stimuli, from neardamaging to overtly destructive of the skin.
The afferent fibers for mechanical nociceptors
have bare nerve endings and, because they are
myelinated, are the fastest-conducting nociceptive
afferents.

PAIN nocireceptors

Pressure on the cell's receptive field with a blunttipped probe elicits no response even if the skin is
indented by 2 mm (A), but the tip of a needle that
punctures the skin produces a clear response (B).
The bottom traces in
parts A and B are
the output of a force
transducer coupled
to the stimulator.
Pinching the skin
with serrated forceps
(C), which is more
traumatic than a pin
prick, produces the
strongest response.

PAIN nocireceptors

The second type is the thermal nociceptors,

are faster-conducting A nociceptors which respond
to the above stimuli as well as to thermal stimuli.
Thermal nociceptors are excited by extremes of
temperature as well as by strong mechanical stimuli.
One group of thermal nociceptors is excited by noxious
heat (temperatures above 45C). A second group
responds to noxious cold (cooling the skin below 5C).
The third type is chemical nociceptors, which
respond only to chemical substances.
A fourth type is known as polymodal
nociceptors, specifically associated with C fibers, other
unmyelinated nociceptors, which respond to high
intensity stimuli such as mechanical, thermal and to
chemical substances like the previous three types.

Silent Nociceptors.
In the skin and deep tissues there are additional
nociceptors called "silent" or "sleep" nociceptors.
These receptors are normally unresponsive to noxious
mechanical stimulation, but become awakened
(responsive) to mechanical stimulation during
inflammation and after tissue injury.
One possible explanation of the "awakening"
phenomenon is that continuous stimulation from the
damaged tissue reduces the threshold of these
nociceptors and causes them to begin to respond.
Many visceral nociceptors are silent nociceptors.

Joint Nociceptors.
The joint capsules and ligaments contain highthreshold mechanoreceptors, polymodal nociceptors,
and "silent" nociceptors.
Many of the fibers innervating these endings in
the joint capsule contain neuropeptides, such as
substance P (SP) and calcitonin gene-related peptide
(CGRP).
Liberation of such peptides is believed to play a
role in the development of inflammatory arthritis.

Visceral Nociceptors.
Visceral organs contain mechanical pressure,
temperature, chemical and silent nociceptors.
The visceral nociceptors are scattered, with
several millimeters between them, and in some organs,
there are several centimeters between each nociceptor.
Many of the visceral
nociceptors are silent. The
noxious information from
visceral organs and skin
are carried to the CNS in
different pathways.

Pain perception
The receptive fields of all pain-sensitive
neurons are relatively large, particularly at the level of
the thalamus and cortex, presumably because the
detection of pain is more important than its precise
localization.
In general, two categories of pain perception
have been described: a sharp first pain and a more
delayed, diffuse, and longer-lasting sensation that is
generally called second pain.
A fibers are responsible for first pain
and C fibers are responsible for the
duller, longer-lasting second pain.

Pain perception

(A) First and second pain, are carried by different axons,

as can be shown by (B) the selective blockade of the
more rapidly conducting myelinated axons that carry the
sensation of first pain, or (C) blockade of the more
slowly conducting C fibers that carry the sensation of
second pain.

Pain perception
In the painful stimulus range, the axons of
thermoreceptors fire action potentials at the same rate
as at lower temperatures.

The number and frequency

of action potential discharge
in the nociceptive axon,
however,
continues
to
increase.

(45C
is
the
threshold for pain).

approximate

Transduction of Nociceptive Signals

Given the variety of stimuli (mechanical, thermal, and
chemical) that can give rise to painful sensations, the
transduction of nociceptive signals is a complex task.
The specific receptors associated with nociceptive
afferent endings are sensitive to both heat and to
capsaicin, the ingredient in chili peppers that is
responsible for the familiar tingling or burning sensation
produced by spicy foods. Since the same receptor is
responsive to heat as well as capsaicin, it is not
surprising that chili peppers seem hot.

(A) Some popular peppers that contain capsaicin.

(B) The chemical structure of capsaicin.

Transduction of Nociceptive Signals

When applied to the mucus membranes of the oral

cavity, capsaicin acts as an irritant, producing protective
reactions. When injected into skin, it produces a burning
pain and elicits hyperalgesia to thermal and mechanical
stimuli. Repeated applications of capsaicin also desensitize
pain fibers and prevent neuromodulators such as substance
P, VIP, and somatostatin from being released by peripheral
and central nerve terminals.
Consequently, capsaicin is used clinically as an
analgesic and anti-inflammatory agent; it is usually applied
topically in a cream (0.075%) to relieve the pain associated
with arthritis, postherpetic neuralgia, mastectomy, and
trigeminal neuralgia. Thus, this remarkable chemical
irritant not only gives gustatory pleasure on an
enormous scale, but is also a useful pain reliever!

Transduction of Nociceptive Signals

Except for chemical sensitivity, nociception could

simply be considered an extreme version of touch and
temperature sensation.
As is so clearly demonstrated in other sensory
systems (e.g., vision, olfaction, and taste), the cloning
and functional characterization of sensory receptors
provides the definitive molecular tools to elucidate the
logic of stimulus detection and perception (Julius &
Nathans 2012).
Indeed, the study
of transient receptor
potential
(TRP)
channels validated the
existence
of
the
nociceptor.

Transduction of Nociceptive Signals

(TRP) channels consist of a large family of ion channels that play

a wide diversity of physiological functions. In mammals, these
family consists of 28 different TRP members grouped in 7
subfamilies. Expressed in a large number of tissues from nerve to
epithelial cells, genetic studies have linked mutations in these ion
channels to human diseases.
Structurally, TRP channels resemble
voltage-gated
potassium
or
cyclic
nucleotide-gated
channels,
having
six
transmembrane domains with a pore between
domains 5 and 6. The majority of TRP channels
are non-selective cation channels that permeate
Na+ and K+, and most of them with significant
Ca2+ selectivity.
Under resting conditions the pore of the channel is closed.
In the open, activated state, these receptors allow an influx of
sodium and calcium that initiates the generation of action
potentials in the nociceptive fibers.

Transduction of Nociceptive Signals

Thermosensory channels, also named thermoTRPs,

define a subfamily of the TRP channels that are activated by
changes in the environmental temperature, from noxious
cold (<15 C) to injurious heat (>42 C).
ThermoTRPs
have become pivotal
drug targets, and the
development
of
therapeutic
compounds
for
pharmacological
intervention
is
actively pursued by
the academy and the
industry.
http://www.brauchilab.org/research.html

Transduction of Nociceptive Signals

Pungent irritants from pepper, mint, and mustard

plants have served as powerful pharmacological tools
for identifying molecules and mechanisms underlying
this initial step of pain sensation.

These natural products have revealed three members of

the transient receptor potential (TRP) ion channel
familyTRPV1, TRPM8, and TRPA1as molecular
detectors of thermal and chemical stimuli that activate
sensory neurons to produce acute or persistent pain.

Transduction of Nociceptive Signals

TRP Family Proteins Involved in Thermal Transduction.

Receptor Protein
TRPV1
TRPV2
TRPV3
TRPV4
TRPM8
TRPA1

Threshold or
Temperature
Range for
Activation (C)
>42
>52
34-38
27-34
<25
<18

Other Characteristics
Activated by capsaicin
Activated by camphor
Activated by menthol
Activated by mustard oil

Temperature dependence of
firing
rates
in
different
thermosensitive
afferents.
Below the firing curves are
shown the ranges over which
the different TRP channels are
activated.

The fourth letter in the name

identifies the subfamily and was
chosen because of the first
member
of
the
subfamily
identified:
V,
vanilloid;
M,
melastatin; A, ankyrin-like.

Transduction of Nociceptive Signals

The so-called vanilloid receptor (VR-1 or TRPV1) is

found in C and A fibers and is activated by moderate heat
(45Ca temperature that is perceived as uncomfortable) as
well as by capsaicin.
Schematic of the VR1/capsaicin
receptor
channel. TRPV1 domains
that confer sensitivity to
various stimuli: capsaicin
(chili pepper) and related
vanilloid
ligands,
extracellular
protons,
(lemon), or peptide toxins
from tarantula (spider).
Another type of receptor (vanilloid-like receptor, VRL-1
or TRPV2) has a higher threshold response to heat (52C), is not
sensitive to capsaicin, and is found in A fibers.

Sensitization
Following a painful stimulus associated with tissue
damage (e.g., cuts, scrapes, and bruises), stimuli in the
area of the injury and the surrounding region that would
ordinarily be perceived as slightly painful are perceived
as significantly more so, a phenomenon referred to as
hyperalgesia. A good example of hyperalgesia is the
increased sensitivity to temperature that occurs after a
sunburn.
This effect is due to changes
in neuronal sensitivity that occur
at the level of peripheral receptors
as well as their central targets.

Sensitization

Peripheral sensitization results from the interaction of

nociceptors with the inflammatory soup of substances
released when tissue is damaged. These products of tissue
damage include extracellular protons, arachidonic acid and
other lipid metabolites, bradykinin, histamine, serotonin,
prostaglandins, nucleotides, and nerve growth factor (NGF),
all of which can interact with receptors or ion channels of
nociceptive fibers, augmenting their respons.

Substances released by damaged

tissues augment the response of
nociceptive
fibers.
In
addition,
electrical activation of nociceptors
causes the release of peptides and
neurotransmitters
that
further
contribute
to
the
inflammatory
response.

Sensitization

The presumed purpose of the complex chemical

signaling arising from local damage is not only to protect the
injured area (as a result of the painful perceptions produced
by ordinary stimuli close to the site of damage), but also to
promote healing and guard against infection by means of
local effects such as increased blood flow and the migration
of white blood cells to the site.
Obviously the identification of the components of the
inflammatory soup and their mechanisms of action is a fertile
area to explore for potential analgesics (i.e., compounds
that reduce pain intensity). For example, so-called
nonsteroidal anti-inflammatory drugs (NSAIDs), which
include aspirin and ibuprofen, act by inhibiting
cyclooxygenase (COX), an enzyme important in the
biosynthesis of prostaglandins.

Pain has been classified into three major types:

1. Pricking pain. Pain caused by a needle, pin prick,
skin cut, etc. - elicits a sharp, pricking quality, stinging
pain sensation carried fast by the A delta fibers.
- is precisely localized and of short duration.
- is also called fast pain, first pain or sensory pain.
- is present in all individuals and is a useful and
necessary component of our sensory repertoire; without
this type of protective pain sensation, everyday life
would be difficult.
- arises mainly from the skin, and carried mainly by A
delta fibers which permits discrimination (i.e., permits
the subject to localize the pain).

Pain has been classified into three major types:

2. Burning pain or soreness pain. Pain caused by

inflammation, burned skin, etc., is carried by the C
fibers (slowly conducted pain nerve fibers).
-is a more diffuse, slower to onset, and longer in
duration.
- it is an annoying pain and intolerable pain, which is not
distinctly localized.
-like pricking pain, burning pain arises mainly from the
skin.
-it is carried by the paleospinothalamic tract. (The old
primitive transmission system for diffuse pain which
does not permit exact localization.)

Pain has been classified into three major types:

3. Aching pain is a sore pain.
-This pain arises mainly from the viscera and somatic
deep structures.
-Aching pain is not distinctly localized and is an
annoying and intolerable pain.
-Aching pain is carried by the C fibers from the deep
structures to the spinal cord

Placebo effect
The placebo effect is defined as a physiological
response
following
the
administration
of
a
pharmacologically inert remedy. The word placebo
means I will please, and the placebo effect has a long
history of use (and abuse) in medicine.

Placebo effect

A common misunderstanding about the placebo effect is

the view that patients who respond to a therapeutically
meaningless reagent are not suffering real pain, but only
imagining it. Although the mechanisms by which the brain affects
the perception of pain are only beginning to be understood, the
effect is neither magical nor a sign of a suggestible intellect. In
short, the placebo effect is quite real.
Understanding the central modulation of pain perception
(on which the placebo effect is presumably based) was greatly
advanced by the finding that electrical or pharmacological
stimulation of certain regions of the midbrain produces relief of
pain. Precisely how pain is modulated is being explored in many
laboratories at present, motivated by the tremendous clinical (and
economic) benefits that would accrue from still deeper knowledge
of the pain system and its molecular underpinnings.

SENSORY INTEGRATION AND SENSORY INTEGRATION DYSFUNCTION

THE TACTILE SENSE

We are always actively touching or passively
being touched by something - other people, furniture,
clothes, spoons. Even if we are stark naked, our feet
still touch the ground, and the air touches our skin.
Therefore, the tactile sense, or sense of touch, is
a huge sensory system that gives us information
needed not only for visual perception, motor planning,
and body awareness, but also for academic learning,
emotional security, and social skills.
Two components make up the tactile sense:
First is the protective (or defensive) system;
Second is the discriminative system.

SENSORY INTEGRATION AND SENSORY INTEGRATION DYSFUNCTION

The Protective/Defensive System

Whether we call it protective or defensive, it's the
same: the "uh, oh!" system. All creatures are born
with a protective/defensive system. Its purpose is to
alert us to potentially harmful stimuli. We need it to
survive.
The receptors for the protective system are in the
skin. Light touch is the stimulus that causes the
receptors to respond.
Sometimes light touch is alarming, such as a
mosquito alighting on our skin. We respond negative,
for self-preservation. For example, when a stranger
gets too close, we shrink: when a lash gets in our eye,
we blink.

SENSORY INTEGRATION AND SENSORY INTEGRATION DYSFUNCTION

The Discriminative System

The second component of the tactile sense is the

discriminative system: the "aha!" system. It tells us:
That we are touching something or that something is
touching us.
Where on our body the touch occurs.
Whether the touch is light or deep.
How to perceive the attributes of the object, such as
its size, shape, temperature, density and texture.

SENSORY INTEGRATION AND SENSORY INTEGRATION DYSFUNCTION

The Discriminative System

It develops as neurological maturation suppresses
the defensive system (the defensive system diminishes
but doesn't disappear. Indeed, messages between the
two systems must continue to flow back and forth all our
lives so we can interpret tactile information
appropriately).
The receptors for this system are in the skin,
especially on the hands and fingers, the soles of the
feet, and the mouth and tongue. Deep touch, or "touch
pressure," is the stimulus that causes the receptors to
respond.

SENSORY INTEGRATION AND SENSORY INTEGRATION DYSFUNCTION

CHARACTERISTICS OF TACTILE DYSFUNCTION

An individual with an inefficient tactile system

may have one or more problems with the integration of
touch sensation. The person may:
1. Be defensive to touch (hypersensitive).
2. Be under-responsive to touch (hyposensitive).
3. Have poor tactile discrimination.

SENSORY INTEGRATION AND SENSORY INTEGRATION DYSFUNCTION

Tactile dysfunction is the inefficient processing in

the central nervous system of sensations perceived
through the skin.
The person who is hypersensitive to touch has
tactile defensiveness, the tendency to react negatively
and emotionally to unexpected, light touch sensations.
The person will react not only to actual touch but also to
the anticipation of being touched. Perceiving most touch
sensations to be uncomfortable or scary, he overreacts
with a fight-or-flight response. A person may typically
avoid unexpected, light touch, but accept, firm touch
(deep pressure), like a bear hug. This person needs
touch information more than a person with a wellregulated tactile sense.

SENSORY INTEGRATION AND SENSORY INTEGRATION DYSFUNCTION

The person who is hyposensitive to touch tends

to under-react to tactile experiences.
Needing extra stimulation, he may constantly
touch objects and people. He may be under-responsive
to touch, whether the touch is soothing or painful.
Unlike the hypersensitive person, who over-reacts
to protect himself', the hyposensitive person may not
react to touch effectively enough to do a good job of selfprotection. In fact, he may he unaware of touch
altogether, unless the touch is very intense.
It is important to understand that the out-of-sync
person may be both hyposensitive and hypersensitive.
For instance, he may jump when someone grazes his
elbow, yet be indifferent to a broken collarbone.

SENSORY INTEGRATION AND SENSORY INTEGRATION DYSFUNCTION

The
person
with
hypersensitivity
(tactile
defensiveness) may:
- Overreact to physically painful experiences, making a
"big deal" over a minor scrape or a splinter. He may be
a hypochondriac.
- React similarly to dissimilar touch sensations. A
raindrop on his skin may cause as adverse a reaction
as a thorn.
- Avoid touching certain textures or surfaces, like some
fabrics, blankets, rugs, or stuffed animals.
- Be unusually fastidious, hurrying to wash a tiny bit of
dirt off his hands.
- Avoid walking barefoot on grass or sand, or wading in
water, or walking on tiptoe to minimize contact with the
ground.

SENSORY INTEGRATION AND SENSORY INTEGRATION DYSFUNCTION

The
person
with
hyposensitivity
responsiveness to touch) may:

(under-

- Seem unaware of touch unless it is very intense.

- Show little or no reaction to pain from scrapes,
bruises, cuts, or shots.
- Hurt other people or pets during play, seemingly
without remorse, but actually not comprehending the
pain that others feel.
- Fail to realize he has dropped something.

SENSORY INTEGRATION AND SENSORY INTEGRATION DYSFUNCTION

The person with poor tactile discrimination may:

- Seem out of touch with his hands, as if they were
unfamiliar appendages.
- Be unable to identify which body parts have been
touched without looking.
- Be fearful in the dark.
- Be unable to perform certain motor tasks without visual
cues, such as zipping, buttoning, and unbuttoning
clothes.
- Have difficulty holding and using tools, such as pens,
scissors, and forks.

CHARACTERISTICS OF PROPRIOCEPTIVE DYSFUNCTION

Proprioceptive dysfunction is the inefficient

processing of sensations perceived through the
muscles, joints, ligaments, tendons, and connective
tissue.
Proprioceptive dysfunction is usually accompanied
by problems with the vestibular and/or vestibular
systems.
Whereas it is common for an individual to have
only tactile, or only vestibular, problems, it is less likely
for an individual to have only proprioceptive problems.

CHARACTERISTICS OF PROPRIOCEPTIVE DYSFUNCTION

The person with proprioceptive dysfunction may:

- Deliberately "bump and crash" into objects in the
environment, e.g., jump from high places.
- Stamp or slap his feet on the ground when walking.
- Kick his heels against the floor or chair.
- Bang a stick or other object on a wall or fence while
walking.
- Rub his hands on tables, bite or suck on his fingers, or
crack his knuckles.
- Like to be tightly wrapped in a blanket or tucked in tight
at bedtime.
- Chew constantly on objects like shirt collars and cuffs,
pencils, and gum.

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