3. Somatosensory system:
function3
Receptors respond to mechanical, thermal & chemical
stimuli within three broad categories:
Proprioceptive (lecture 6,10)
Body and limb position
Enteroreceptive (KIN 305)
Internal state of the body
Exteroreceptive
Touch
Temperature
Pain (nociception)
4. Somatosensory fibers
4
Afferent fibers
carrying info of
different
modalities are of
different sizes
What property of large
diameter axons allows
for increased speed of
conduction?
5. Somatosensory transduction
5
two classes: simple or complex
mechanical deformation, heat or
chemical stimulus within
receptive field opens ion
channels
causes a local depolarization
= receptor potential
propagated by electrotonic
conduction to axon hillock
Stimulus intensity coded by
number of receptors activated,
and frequency of AP
Receptor potential
6. Somatosensory transduction
6
How to convey modality of sensation?
Labelled line coding
Type of sensation felt when receptor is stimulated
determined by where the fiber synapses in the CNS
Examples of “fooled” senses:
Chewing minty gum activates cold sensitive
thermoreceptors sensation of cold
Seeing stars after being hit on head forceful blow
activates photoreceptors see spots
11. Cutaneous mechanoreceptors:
summary11
Receptor Sensation Adaptation rate Receptive field
Free nerve endings Itch, tickle, pain Tonic or phasic Large or small
Ruffini endings Stretching of skin,
deep pressure
Tonic Large
Merkel discs Fine touch and
pressure
Tonic Small
Meissner corpuscle Fine touch, pressure,
slow vibration
Phasic – moderate Small
Hair follicle Crude touch,
movement of hairs
Phasic – moderate Small
Krause bulbs Fast vibration Phasic - fast Small
Pacinian corpuscle Pressure, fast
vibration, tickling
Phasic - fastest Large
12. Thermoreceptors
12
Free nerve endings with high thermal sensitivity
Temperature change activates family of ion channels on the
receptor membrane = TRP (transient receptor potential)
channels
Each TRP channel has a unique temperature threshold of firing,
and is sensitive to various chemical agonists
13. Thermoreceptors: tonic
responses13
Guyton Fig. 48-10
Warmth receptors:
Narrow temperature range
Begin firing at 30ºC, rise
steeply with increased
temperature, then stop
abruptly
Sensation of pain begins
>45ºC
Cold receptors:
Broader temperature range
Maintain steady discharge
rates, with increased firing
20-30ºC
<15ºC neuronal firing
ceases
Tonic response of thermoreceptors and
thermosensitive pain fibers
14. Thermoreceptors: phasic
responses14
Kandel Fig. 22-9
Thermoreceptors are more responsive
to changes in temperature than to
constant temperature
A phasic response in both warm and
cold receptors occurs when the
temperature is changed
Thermoreceptors adapt to a new
steady state firing level is stimulus
is maintained
15. Nociceptors
15
Free nerve endings that respond to intense stimuli
Types:
Mechanical
Strong pressure, sharp objects
Thermal
Burning heat (>45ºC)
Noxious cold (variable)
Chemical
pH extremes
Environmental irritants
Internal neuroactive substances
Polymodal
Sensations mediated by Aδ fibers (sharp, intense pain) and C fibers
(persistent, dull pain).
Check: Which fibers
are myelinated?
16. Nociceptors: hyperalgesia of
inflammation16
Nociceptors are non adapting receptors
Primary hyperalgesia: damaged tissue
has increased sensitivity to pain
Reduced threshold to pain
If normally non painful stimuli felt as
painful: allodynia
Increased intensity of sensation
Spontaneous pain
Inflammatory response releases
bradykinin, prostaglandins, serotonin,
substance P, K+
, H+
Substance P activates mast cells
release histamine activates
nociceptors
B&B Fig. 13-26
17. Somatosensory projections:
dermatomes17
Sensory neurons (dorsal
root ganglion cells) enter
the spinal cord through the
dorsal roots
Each dorsal root innervates
a field of skin called a
dermatome
Dermatomal map used
to determine level of
lesion of spinal injury
Epidural analgesia
blocks sensations thru
out several dermatomes
B&L Fig. 7-4
19. 1. Dorsal column-medial lemniscus
pathway19
Carry signals from mechanoreceptors of the skin, joints
& muscle fine, discriminatory touch
Information has high spatial & temporal resolution
Large, myelinated fibers with rapid conduction velocities
1º
afferents terminate & form synaptic connections with
2nd
order neurons in the dorsal column nuclei within the
medulla
2nd
order neurons cross over at the medulla and continue
to the thalamus via the medial lemniscus pathway
After thalamic processing, 3rd
order neurons project to
the primary somatosensory cortex
20. 2. Anterolateral pathway
20
Pain, temperature, crude touch, tickle, itch, sexual
sensations
Low spatial or temporal resolution
Small, myelinated/unmyelinated fibers with slower
conduction velocities
1° afferents terminate upon entering spinal cord &
synapse on 2nd
order neurons
2nd
order neurons cross to contralateral side, ascend to
brain via anterolateral quadrant in spinal cord & project
to thalamic nuclei
After thalamic processing, 3rd
order neurons project to
the primary somatosensory cortex and other cortical
areas
21. Somatosensory cortex:
somatotopy21
B&B Fig. 14-11
Spatial orientation of signals form
different parts of the body: somatotopy
Size of somatotopic areas is proportional
to density of sensory receptors in that
body region
Map is plastic (modifiable)
size of cortical region representing
particular portion of body surface can
expand or contract depending on use
of that body region
Pain and temperature localization not as
precise
Integration probably happens more in
the reticular formation and thalamus
22. Pain sensation: referred pain
22
Pain from visceral nociceptors is poorly localized, can be felt as
pain on surface areas
Knowledge of referred pain maps important in clinical diagnosis
Somatic and visceral afferents
may converge on same 2nd
order
neuron
Guyton Fig. 48-6Kandel Fig. 24-3
23. Pain sensation
23
Sensation of pain intensity not necessarily linked to activation of nociceptors;
under CNS control
Perception of pain and pain tolerance is subjective
Gate control theory of pain
Activation of non-painful fibers (Aα) sends inhibitory signals to
nociceptive afferents traveling to the CNS
Mechanism of acupressure analgesia?
Phantom limb pain
Pain felt even though nociceptors no longer present in missing limb
Peripheral sensitization
Somatosensory reorganization
Neuropathic pain
Damage to Aδ or C fibers may increase sensitivity or cause
spontaneous AP firing
24. Pain disorders: CRPS
24
Complex regional pain syndrome (CRPS)
Neuropathic pain disorder involving peripheral
and central mechanisms (autonomic nervous
system)
Changes in somatosensory systems
processing thermal, tactile, noxious stimuli
Local edema, altered sweating, redness,
skin temperature changes, burning pain,
hyperalgesia, allodynia
Acute: warm, red extremities
Chronic: cool, bluish extremities
CRPS I – no documented nerve injury
CRPS II – presence of nerve injury
Surgery, fracture, crush injury, sprains, but can
develop even after minimal injury
WebCT readings: Complex Pain Syndrome
Left arm affected by CRPS
Right foot affected by CRPS
25. Pain disorders: CRPS
pathophysiology25
Peripheral and central sensitization
Tissue injury release of substance P and
bradykinin increased excitability of
nociceptive neurons in periphery and spinal
cord
Increased local, systemic, and CSF
inflammatory factors
Reduced density of Aδ and C fibers
Altered SNS function and sympatho-afferent
coupling
Expression of adrenergic receptors on
nociceptors
Symptoms worsened by emotional arousal
Reduced representation of affected limb in
somatosensory cortex
Genetic predisposition
Bruehl S. Anesthesiology 2010WebCT readings: Complex Pain Syndrome
26. Objectives
After this lecture you should be able to:
List the structure and function of the various cutaneous
mechanoreceptors
Describe the mechanism of somatosensory transduction,
including modality coding and receptor adaptation
Differentiate between tonic and phasic responses of
thermoreceptors
Compare and contrast the function and anatomy of the dorsal
column-medial lemniscus pathway with the anterolateral
pathway
List the factors affecting pain sensation
Relate what we’ve learned in this course so far to the different
theories of CRPS pathophysiology
26
27. 27
1. Predict on which side of the body and what sensations
would be impaired if there was a spinal cord
transection at T4 of:
a) The left dorsal column-medial lemniscus pathway
b) The left anterolateral pathway
2. Describe what would happen to both cold and warm
thermoreceptors in response to an increase in
temperature from 35ºC to 40ºC , sustained for 5 min,
and removed.
3. Meissners corpuscles are __________________
adapting receptors with __________________ receptive
fields.
Test your knowledge
Cutaneous mechanoreceptors because we already talked about mechanically gated receptors: Hair cells in vestibular and auditory system
System of sensory receptors and central projections that transduce, encode, and ultimately perceive information generated by stimuli arising from both the external and internal environment
Next lecture, muscle proprioceptors and vestibular system
Differences between somatosensory system and special senses is that somatosensory receptors are diffusely located thru out the body instead of densely distributed to a specific organ. Special senses convey information to the brain via single nerve bundle whereas somatosensory information arrives via the spinal cord dorsal roots and cranial nerves
Enteroreceptive – mechanoreceptors detecting distension of gut and fullness of bladder
. Although the basic senses—somatic sensation, vision, audition, vestibular sensation, and the chemical senses—are very different from one another, a few fundamental rules govern the way the nervous system deals with each of these diverse modalities. Highly specialized nerve cells called receptors convert the energy associated with mechanical forces, light, sound waves, odorant molecules, or ingested chemicals into neural signals that convey information about the stimulus to the brain. These afferent sensory signals activate central neurons capable of representing both the qualitative and quantitative aspects of the stimulus (what it is and how strong it is), and in some modalities—somatic sensation, vision, and audition—the location of the stimulus in space (where it is).
The clinical evaluation of patients routinely requires an assessment of the sensory systems to infer the nature and location of potential neurological problems. Knowledge of where and how the different sensory modalities are transduced, relayed, represented, and further processed to generate appropriate behavioral responses is therefore essential to understanding and treating a wide variety of diseases. Accordingly, these chapters on the neurobiology of sensation also serve to introduce some of the major structure/function relationships in the sensory components of the nervous system.
Increased length constant due to decrease axoplasmic resistance
The local depolarization produced is called a generator or receptor potential.
Receptor potential increases with stimulus strength.
Site of AP initiation = trigger zone
Stimulus intensity encoded in:
firing rate of neuron (number of action potentials per second) or
number of activated sensory receptors
The firing rate of single receptor is frequently an exponential function of stimulus intensity.
Ruffini&apos;s end organs detect tension deep in the skin.
Meissner&apos;s corpuscles detect changes in texture (vibrations around 50 Hz) and adapt rapidly.
Pacinian corpuscles detect rapid vibrations (about 200–300 Hz).
Merkel&apos;s discs detect sustained touch and pressure.
Mechanoreceiving free nerve endings detect touch, pressure and stretching
Hair follicle receptors are located in hair follicles and sense position changes of hairs.
Krause bulbs – innervate border areas of dry skin and mucous membranes (ie. lips external genitalia)
Pacinian Corpuscle – glaborous and hair skin. vibration at 200-300Hz due to specialized capsule
Repetitive discharge = train of action potentials
Slowly adapting mechanoreceptors continue responding to a stimulus, whereas rapidly adapting receptors respond only at the onset (and often the offset) of stimulation. These functional differences allow the mechanoreceptors to provide information about both the static (via slowly adapting receptors) and dynamic (via rapidly adapting receptors) qualities of a stimulus.
A receptive field is a region that causes activation of a sensory neuron when stimulated.
Stimulus may not be perceived unless several sensory receptors are activated (spatial summation) or unless the firing rate is sufficiently high (temporal summation) to bring postsynaptic neurons to threshold.
Small receptive fields are characteristic of Meissner’s corpuscles and Merkel’s disks (2-4 mm in diameter).
Pacini’s and Ruffini’s corpuscles have large receptive fields.
Example: Ruffini’s corpuscles respond to stretch of skin even at some distance from receptor site.
Meissner’s Corpuscles (FA1 afferents)
also rapidly adapting receptors
respond best to vibration in 30-40 Hz range
Meissner’s corpuscles are located in ridges of the dermis of glabrous skin; they are much smaller than Pacini’s corpuscles.
Meissner’s corpuscles receive innervation from 2-5 axons, while one axon innervates about 20 corpuscles; other receptors innervated by single axon.
The rapidly adapting receptors (Pacini’s and Meissner’s corpuscles) provide contact information, slip information, &contrast information (e.g. important for edge detection).
Merkel’s Disks (SA1 afferents)
slowly adapting receptors that
respond to touch
Merkel’s disks are located at border of dermis and epidermis of glabrous skin. Merkel cells seem to best encode spatial characteristics of textured surface.
Pacinian Corpuscles (FA2 Afferents)
respond transiently to applied pressure
during sustained pressure layers slip reducing deformation of receptor membrane
also respond during restitution of membrane when pressure is released
respond best to vibration in 200-300 Hz range
Ruffini’s Corpuscles (SA2 afferents)
slowly adapting receptors
respond to indentation of the skin
(touch & low frequency vibration)
Hair Follicle Receptors
innervated by free nerve endings that wrap around or run parallel to follicle
bending of hair causes deformation of follicle which deforms nerve endings and modulates their firing frequency
Meissner’s Corpuscles (FA1 afferents)
also rapidly adapting receptors
respond best to vibration in 30-40 Hz range
Meissner’s corpuscles are located in ridges of the dermis of glabrous skin; they are much smaller than Pacini’s corpuscles.
Meissner’s corpuscles receive innervation from 2-5 axons, while one axon innervates about 20 corpuscles; other receptors innervated by single axon.
The rapidly adapting receptors (Pacini’s and Meissner’s corpuscles) provide contact information, slip information, &contrast information (e.g. important for edge detection).
Merkel’s Disks (SA1 afferents)
slowly adapting receptors that
respond to touch
Merkel’s disks are located at border of dermis and epidermis of glabrous skin. Merkel cells seem to best encode spatial characteristics of textured surface.
Pacinian Corpuscles (FA2 Afferents)
respond transiently to applied pressure
during sustained pressure layers slip reducing deformation of receptor membrane
also respond during restitution of membrane when pressure is released
respond best to vibration in 200-300 Hz range
Ruffini’s Corpuscles (SA2 afferents)
slowly adapting receptors
respond to indentation of the skin
(touch & low frequency vibration)
Hair Follicle Receptors
innervated by free nerve endings that wrap around or run parallel to follicle
bending of hair causes deformation of follicle which deforms nerve endings and modulates their firing frequency
In accordance with this functional diversity, TRP channels are implicated in a multitude of physiological processes, ranging from Ca2+ and Mg2+ homeostasis and regulation of the vascular tone to bone development, taste perception, temperature sensing and vision.
These membrane receptors belong to a class (family) of proteins called the transient receptor potential proteins; these receptors bind to the chemical capsaicin, the molecule in chili peppers that gives them their heat. Other binds different chemicals including camphor, menthol, and mustard oil
Stimulus-response curve of cold receptors is non-monotonic so temperature cannot be signaled by firing rate alone; perceived temperature is determined by relative activity of cold and warm receptors.
Rapid changes in skin temperature evoke dynamic responses (transient increase in firing rate of warm receptors for increase in temperature or transient increase in firing rate of cold receptors for decrease in temperature).
Neuroactive substances include histamine and bradykinin in sites of infection
Adelta: fast, myelinated
C: slow, unmyelinated
Increased pain following sunburn. Secondary hyperalgesia describes pain that occurs in surrounding undamaged tissue.
large 1 afferent axons bifurcate in dorsal column and branches:
rostrally to dorsal column nuclei (DCN)
caudally, over a distance of several spinal segments
Collateral branches of primary afferents project ventrally into grey matter of spinal cord where they synapse on interneurons and motoneurons.
The person is on all fours because this is how dermatomes develop during development, they are distorted during embryological development.
Epidural from L3-4 blocks sensation in uterus and lower dermatomes. Inserted into epidural space because this is the location of the dorsal root ganglia
Carry signals from mechanoreceptors of the skin, joints & muscle fine, discriminatory touch
Large, myelinated fibers with rapid conduction velocities
Information has high spatial & temporal resolution
1º afferents terminate & form synaptic connections with 2nd order neurons in the dorsal column nuclei within the medulla
2nd order neurons cross over at the medulla and continue to the thalamus via the medial lemniscus pathway
Pain, temperature, crude touch, tickle, itch, sexual sensations
Small, myelinated fibers with slower conduction velocities
Low spatial or temporal resolution
1 afferents terminate upon entering spinal cord & synapse on 2nd order neurons
2nd order neurons cross to contralateral side, ascend to brain in ventral part of lateral funiculus & project to thalamic nuclei
3rd order neurons project to somatosensory cortex & other cortical areas
START!
Mechanoreceptors of the skin, joints and muscle provide tactile, discriminative, and proprioceptive information.
A high spatial and temporal resolution refers to the ability to discriminate details of spatial location and stimulus intensity.
Fasciculus gracilis: fibers conveying information from lower limbs are in the medial region
Fasciculus cuneatus: fibers from upper limbs, trunk and neck are in the lateral region
DCN = gracile and cuneate nuclei
most prominent ascending pathway for pain and thermal sensations, also carries crude tactile sensation
thalamic nuclei include VPL nucleus & intralaminar complex & others
cortical projections of 3rd order neurons are involved in affective responses such as the cingulate gyrus and insula, which have limbic system functions
Somatotopic order in the human primary somatic sensory cortex. (A) Diagram showing the region of the human cortex from which electrical activity is recorded following mechanosensory stimulation of different parts of the body. The patients in the study were undergoing neurosurgical procedures for which such mapping was required. Although modern imaging methods are now refining these classical data, 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) Cartoon of the homunculus constructed on the basis of such mapping. Note that 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. A similar disproportion is apparent in the primary motor cortex, for much the same reasons (see Chapter 17). (After Penfield et al., 1953, and Corsi, 1991.)
after denervation, representation of neighboring skin regions expands to occupy that of denervated region (may enhance sensitivity to stimuli or ability to process)
- signals from neighboring regions to compensate for missing input {Fig. 14-13}
excessive use of one digit at expense of others results in expansion of its cortical representation
Allodynia: different form hyperalgesia in that a normally non painful stimulus, such as light touch, is felt as extrememley painful, (A fiber mediated) whereas in hyperalgasia the painful stimulus intensity is heightened (A or C fiber mediated)
Reduced density not known if it is a cause or a result of CRPS
Left side of body fine discriminatory touch below T4
Right side of the body, pain temp heat below T4
3. Moderately adpating phasic receptors with small receptive fields.