CHAPTER 3: THE SOMATOSENSORY SYSTEM TOUCH, FEELING, AND PAIN

Neural Basis for Somatosensory Perception

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Theory of Receptor Specificity individual receptors and nerve endings are selectively
sensitivity to a particular form of energy impinging on the skin and indicates location of
stimulus
Patter Theory specificity of touch sensations arise through the overall pattern of activity
across a broad spectrum of receptors
Mechanoreceptors senses touch

Mechanoreceptors and Afferent Fibres

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Mechanoreceptors are Modified bipolar neurons: Axons and dendrite fuse as one fiber with
the receptor end as peripheral branch and the spinal cord end as central branch
o Dorsal Root ganglion is the cell body
o Action potentials are produced at the receptor site directly
- Transmission Speed: Larger the diameter and greater the degree of Myelination, the faster
the nerve signal transmission In order of speed A >A >A > C
- Response Adaptation: Refers to how the fibres respond to continuous touch stimulation
o Slowly Adapting (SA) Fires as long as the stimulus is maintained. Low Temporal
resolution
o Fast Adapting (FA) A transient fast neural firing that subsides quickly. High
temporal resolution.
- Receptive Field Size: influenced by the location on the skin and type of receptor
o Type I (SA-1 / FA-1) superficial small receptor field high spatial resolution
Types of
MechanoSkin
Fibr Respons RF Size
Stimulus
Receptors
receptors
Location
e
e Type
Detected
Type
Pacinian
Deep (hairy / hairless)
A
FA-I
Large
Vibrations
Ruffini
Deep (hairy / hairless)
A
SA-II
Large
Steady
Encapsulat
Pressure
ed
Meissner
Superficial (hairy /
A
FA-I
Small
Flutter;
hairless)
Motion
Merkel Disc
Superficial (hairy)
A
SA-I
Small
Steady
Accessory
Pressure
Lanceolate
Superficial (hairy)
Small
Hair movt
Free Nerve Endings
Superficial (hairy)
A
Mixed
Variable
Temp; Pain
o Type II (SA-II / FA-II) deep big receptor field low spatial resolution

Spinal Mechanisms and Signal Transfer

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Spinal Nerves and Ganglia: 31 pairs of spinal nerve emerging left and right of the spinal
cord
o Each spinal nerve is divided into 2 roots dorsal and ventral root and then coming
together again
o Bell-Magendie law basic principle regarding the separation of function of the roots
Dorsal Root carries sensory signals into the spinal cord
Ventral Root carries motor signals out and into the muscles
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o DR Ganglion only present in the dorsal root

Dermatomal Map: Map representing the skin surface devoted to all spinal nerve
o Dermatome discrete areas on the body surface that are represented by a single
nerve
Signal transmission through the Spinal Cord, Relay nuclei and Cortex:
o Dorsal column-medial Lemniscus Pathway: Large Diameter, myelinated fibres
(A, A) tactile + proprioceptive signals
DRG neurons Dorsal horn (1st relay site) synapse with motor neuron in
ventral horn OR
Go up dorsal column of spinal cord directly dorsal column nuclei (2nd
relay site) cross to the other side of the brain ascend as fibre bundle
call medial lemiscus Thalamus (3rd relay site) direct projection to
cortex as fibre bundle call internal capsule
o Anterolateral Pathway: Small diameter, non-myelinated fibres (A, C) pain and
temperature signals
DRG Neurons Dorsal horn (1st relay site) cross over to the other side of
the spinal cord enter the anterolateral system Most terminate in relay
nuclei in medulla, midbrain and thalamus those in thalamus project to
the cortex in a diffuse manner
Parallel Processing and Modality Segregation:
o Modality Segregation presence of two parallel pathways that permits independent
transmission of different types of touch signals.
o Advantage allows the brain to process the 2 types of signals separately leads to
richness and diversity of perceptual experiences
Somatotopic Representation: neighbouring neurons process tactile signals from adjacent
locations on the body surface.

Somatosensory Cortex
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Area S1: Primary Somatosensory cortex; serves as the starting point for cortical processing
of info
Info is processed in serial (increasing order of integration) and parallel (separate signals)
Somatotopic Map: The cortex contains and entire representation of the body surface. In
face each subdivision of S1 also contains the total body representation (Homunculus).
o Contralateral representation the Left cortex represents right body and vice versa
o Non linear representation body parts are represented in proportion to its relative
importance (hands and face are represented more than toe and trunk)
Columnar Organization: Responses of the different types of receptors are organized in
columns run vertically from the cortical surface to the white matter
o Columns of FA and SA are arranged in an alternating fashion. N
o Neurons in each layer respond to the same modality at any given area

Somatomotor Circuits
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Somatomotor Control in Spinal Cord: Motor signals emerge out of the spinal cord through
the ventral roots and proceed through the spinal nerves to innervate the muscles.
o 1st source of innervation descending fibres which carry commands from the cortex
o 2nd source of innervation afferent fibres from the Somatosensory system
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Somatomotor Control in the Cortex: The primary motor cortex and Somatosensory cortex
are in close proximity to allow a more precise registration int eh connectivity.
o The Somatosensory projections to the motor cortex are homotopic, meaning the
motor cortex receives projections from the Somatosensory cortex for the same
location on the body map.

Perceptual Aspects of Tactile Sensation

Intensity and Sensation
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Esthesiometers: Device used to measure the sensitivity of skin

Absolute Threshold: absolute threshold of touch is very small but also depends on the site
on the body
o Facial regions are more sensitive than extremities and lower end of the body.
o Reduction in skin elasticity will reduce sensitivity (i.e. Callus thickened skin less
elastic)
Difference Thresholds: Multiple factors that affects this
o At low or high intensities, the constant in Webers law is actually not constant
o Site on the body skin areas that have low touch threshold more sensitive to
stimulus change lower Webers constant
Sensory magnitudes:
o Single indentation in hairless skin produce sensation magnitudes that increased
linearly with depth of skin displacement
o Skin indentation on hairy skin smaller sensory magnitudes

Spatial Factors
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Two Point Limen: the smallest separation of two points applied simultaneously to the skin
that can still be discriminated
o Compass Test 2 stimuli given simultaneously
o Point Localization error test 2 stimuli given successively
Law of Outward Mobility: 2-point limen improves 20x as one moves fromt eh shoulder
toward the finger tips
Facial region are most sensitive to touch, but fingers have the highest spatial
discrimination.
The feet have good discrimination ability but poor sensitivity.
Mechanoreceptor Properties Related to 2-Point Limen
o Surface receptor with smaller receptive field
o Increase density of the receptors with small receptive field ( SA-I and FAI)
Hyperacuity: spatial discrimination thresholds that are much lower than the traditional
two-point limen

Temporal Factors
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Adaptation: when the perceived intensity of a sensory stimulus is reduced to a point where
it is no longer consciously experienced
o The greater the area of the stimulus, the faster the adaptation
o The greater the intensity or force, the slower the adaptation
3

o Less sensitive areas of the body adapt the quickest

o Adaptation occurs b/c of the lack of stimulus movement
Vibrotactile Stimulation: Vibrating stimulus between 200-300Hz produces the lowest
absolute threshold
o Seems to only work with large receptive field over large area of the skin, thus a
vibrating pin wont reduce threshold EXCEPT if the frequency is much lower
Duplex Theory: 2 separate and independent channels mediate the temporal nature of
touch. Later revised to 4 different channels each having separate properties and each
triggered by specific types of mechanoreceptors.
o A frequency independent channel operate at low frequencies
o Frequency dependent channel operate only at high frequencies
Mechanoreceptor properties related to Vibrotactile Processing
o Meissners corpuscles more sensitive to low frequency stimulation which
correspond to the fact that it has small receptive field
o Pacinian corpuscle sensitive to high frequency stimuli which correspond to the fact
that is has large receptive field
o Merkel receptors produce the sensation of pressure and respond best to static or
near-static stimuli
o Ruffini receptors involved in steady pressure detection, but can provide a buzz like
sensation at high temporal frequencies
Frequency Discrimination and Pitch Perception
o Pitch The perception of frequency. Differences of threshold in pitch is complicated
by 2 factors..
Threshold values actually depend on the frequency
Intensity of stimulus can be affected by small changes in frequency

Thermal Sensations
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the perceived absence of heat or cold sensation occurs only when our body is exposed to
a certain level of heat or thermal energy
Physiological Zero: Range of temperature in which we feel neither warmth nor cold
Thermoreceptors: Divided into warm or cold receptor that are unencapsulated and are
distributed throughout the body. Very sensitive to local change in temperature
o Warm Receptors Slow C fibres; Cold Receptors A fibres (slightly faster)
o Both have small receptive fields
o Slow changes in temp produce little neural activity, but rapid changes are very
effective
o Overall effective in detecting change, rather than absolute temperature
Perceptual Aspects of Temperature: Actual perception depends on location, area and
current body temp
o Temperature change required for detection in the extremities is higher than the
facial region
o Spatial Summation stimulation of larger skin area results in lower threshold values
o The rate of growth of perceived warmth as a function of temperature is inversely
related to area very small stimulation area produce faster rates and vice versa.
Only applies to warm and not cold
o Opponent Process warmth and cold are different dimensions of the same sensory
experience
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Tactile Communication
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Braille System: 64 different combination of raised dots organized into groups of 6 (2x3)
Moon System: Modified structure of the alphabet. More blurring making this system
unpopular, but easy to learn
Tactile Mapping System: provides a pattern of stimulation on the skin that portrays the
spatial information present in visual stimulus
o Optacon Optical-to-tactile convertible Vibrating pin on which the user places his
index finger on the array.
o Tactile Vision Substition System (TVSS) produced a tactile display of images from a
video camera directly onto a vibrotactile belt that was worn around the abdomen.
Communication with Deaf-blind Individuals:
o Sign Language instead of watching they touch their hands
o Tadoma (aka tactile lip learning) person places their hands and fingers over the
speakers lips and face.

Proprioception and Kinesthesis

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Kinesthesis: the ability to feel movements of the limb and body

Proprioceptors: neural elements that reside in muscles, tendons and joints to allow
kinesthesis
o Senses only position and movement of the various body parts

Neural Foundations
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Muscle Spindle Receptor: Slow adapting fibre that is very sensitive to stretch
Golgi Tendon organ: encapsulated structure located at the junction between the muscle
and tendon. Sensitive to change in tension
Both types of nerve fibres are A type large diameter and highly myelinated. Also go
through DRG and synapse with motor neuron for reflexes
Information is reached in the S-1 area in aprticular the 3b section. 3a processes tactile
signals

Perceptual Aspects
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Hip joints appears to be the most sensitive

Decreasing sensitivity knee > ankle > elbow > wrist > finger base
o Measurement is complicated by direction of movt, degree of stretch nad precise
way in which the movt are made
receptors are responsible for seasation of force, pressure and weight through tension
placed on the muscle
Corollary Discharge: a motor copy of the voluntary movt transmitted to the sensory cortex

Active Touch and Haptic Perception

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Haptic Perception: The combination of tactile and kinesthetic sensory experience from
active touch (the physical manipulation of an object)
Active Touch: Ability to identify an object by active touch is far superior to passive touch. It
allows greater interaction and exploration that enhances our ability to gather information
about the object
o It is wilful in nature engages the mind
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The recruitment of proprioceptors that provide kinesthetic information (i.e. finger

following contours)
Exploratory Strategies in Haptic Perception: Lateral motion, pressure, enclosure, and
contour following
o

Loss of Haptic Perception

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Peripheral Neuropathy: sensory loss due to a widespread degeneration of the large

myelinated fibres of the peripheral nervous system.
o Makes all limb movt nearly impossible to coordinate

Nociecption and Pain

Sensory mechanisms of Nociception
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Nociception: The capture of noxious stimuli through nociceptors to provide the brain with
sensory signals
Pain: the perceptual experience that is generated by the brain from incoming nociceptive
signals.
Neural pathways:
o Microneurography compares electric signals captured from individual nerve fibres
with different kidns of sensory stimulation applied to the skin.
Studies show that activities is detected in the smallest fibres A and C fibres.
o 1st synapse with dorsal horn cross over to the opposite side of the spinal cord and
enter in the anterlateral system various projections to various subcortical nuclei
Somatosensory cortex
Referred Pain: Pain that arises in deeper structures of the body is actually felt elsewhere
o A result of convergence of afferent fibres onto the spinal cord from different parts of
the body.
Phantom pain: pain experienced in a limb that no longer exists due to amputation
o Dorsal horn neurons become hyperactive when sensory input is removed.
o Central Pain nociceptive signals are not generated at the peripheralnerve
terminals, but rather at a more central site (i.e. brain or spinal cord)
Gate Control Theory
o Descending influences from the brain as well as signals from large afferents
converge upon the spinal cord and modulate the transmission of incoming
nociceptive signals through the C fibres.
o An Inhibitory gate to inhibit pain signals going up through the projection neuron
inhibitory interneuron
o Central influences from the brain and large afferent fibers synapse on the inhibitory
interneuron increases the inhibitory effect of the interneuron on the projection
neuron reduced pain.
o C fibre and afferent nerves itself will positively stimulate the projection neuron
increased pain

Measurement of Pain
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The Univariate Concept of Pain: the experience of pain is measured in terms of its overall
integrated impression

One difficulty in applying the standard threshold: the subject has to make a decision
as to the quality of that sensation and whether or not it rises to the definition of
pain.
o Measurement: Use ascending Method of Limits to make to find detection threshold
and pain threshold
Or use a scale or descriptive words
The Multivariate Concept of Pain: assessing and examining the different qualities of pain
o

Control of Pain
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Pharmacological Approach: Using Analgesics

o Anti-inflammatory drugs aspirin, Tylenol, ibuprofen (works at local level)
o Antidepressants, anticonvulsants, and muscle relaxants
o Opioid Analgesics (most powerful) codeine and morphine (works in brain)
Endogenous opiates = endorphins produced by the brain
Anesthetic, Surgical and Neurostimulatory Appraches:
o Anesthetics Local level or in spinal cord to stop transmission of pain signals (i.e.
epidural procedure)
o Surgical Peripheral nerves to spinal cord (cordotomy); frontal lobe (lobotomy).
o Electrical Stimulation active suppression of pain by blocking neural transmission,
endorphin production, descending signals to inhibitory interneurons
TENS electrical stimulation of the peripheral nerves
Psychological Approaches: Behavioural therapy relaxation training, distraction
techniques and hypnosis

Lecture Notes
Tactile Psychophysics:
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What
o
o
o
o

can we measure:
Absolute threshold
Difference threshold
Two-point limen sensation is from 1 point or 2 points?
Point localization error stimulate 1 point successively to see if they can notice the
space difference
Vibration as a function of frequency

Perception and Action:

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Perception for action

o using vision to locate the coffee cup
o using somatosensation to grasp and move it towards our mouth
o using audition to orient towards your housemates
o using taste to prevent swallowing the rotten cream
Action for perception
o saccadic eye movements keep an object on the retina stable
o use of your hands to extend the pinnae to capture faint sound
o in somatosensation, action plays an even larger role: exploratory procedures to
gather information about objects and surfaces - haptics

Haptic Perception: knowledge of the world that is derived from sensory recep[tors in skin,
muscles, tendons, and joints usually involving active touch as a method of exploration
Somatosensory illusions: ownership through consistency
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Rubber band illusion

Disappearing hand trick

Somatosensory response to Tickling

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Self-induced tactile stimulation

Self-induced movement, no tactile stimulation
Externally induced tactile stimulation