The somatosensory system

(touch & proprioception)

Subject

Status Completed

Week 4

Property Chapter 9

Afferent Fibers Convey Somatosensory Information to the Central Nervous

System

The cell bodies of afferent fibers reside in a series of ganglia → lie alongside the
spinal cord & brainstem ~ considered part of the peripheral nervous system.

Action potentials generated in afferent fibers by events - occur in the skin, muscles/
joints → propagate along the fibers ~ past the locations of the cell bodies in the
ganglia until reaching a variety of targets in CNS.

Pseudounipolar - neurons in the dorsal root ganglia.

Conduction of electrical act. thru. the membrane of cell body - not an obligatory step
in conveying sensory info. to central targets.

Cell bodies of sensory afferents - critical role in maintaining the cellular

machinery → mediates transduction, conduction, transmission by sensory afferent
fibers.

Sensory transduction - the process of converting the energy of stimulus into

electrical signal.

a stimulus alters the permeability of cations channels in the afferent nerve

endings → generate a depolarising current (receptor potential).
→ rate of action potential firing - proportional to the magnitude of the
depolarisation.

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 Piezo channels - have more than 30 transmembrane domains.

Purified Piezo proteins - reconstituted in artificial lipid bilayers → form ion

channels ~ transduce tension in the surrounding membrane.

Piezo2 - expressed in subsets of sensory afferent neurons as well as in other

cells.

Afferent fibers terminals - detect and transmit touch sensory stimuli

(mechanoreceptors)
→ encapsulated by spec. receptor cells ~ help tune the afferent fibers to part.
features of somatic stimulation.

lower threshold for action potentials generation ~ more sensitive to sensory

stimulation.

Free nerve endings - afferent fibers that lack spec. receptor cells.
→ imp. in the sensation of pain.

Somatosensory Afferents Convey Different Functional Information

Somatosensory afferents differ → define distinct classes of afferent ~ ea. makes

unique contributions to somatic sensation.

Afferent Ia - supply the sensory receptors in the muscles. (large diameter)

Ab afferents- convey info. subserving touch. (small diameter)

As and C afferents - convey info. about pain & temperature. (very small diameter)

⇒ determines the action potential conduction speed - matched to properties of the

central circuits + various behaviour demands for employed sensory afferent.

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 Receptive field - area of the skin surface over which stimulation → results in a sig.
change in the rate of action potentials.

A given region of body surface - served by sensory afferent → vary sig. in the size
of their receptive fields.

There are systematic regional variations in the avg. size of afferent RF → reflect the
density of afferent fibers supplying the area.

dense innervation - small region area (fingers, lips, toes)

⇒ regional diff. in RF size & innervation density - major factors that limit the spatial
accuracy w. tactile stimuli can be sensed.

Two-point discrimination - min. interstimulus distance required → perceive 2

simultaneously applied stimuli as distinct ~ vary across the skin surface.

Sensory afferents - differentiated by the temporal dynamics of their response to

sensory stimulation.

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 Some afferents fire rapidly when a stimulus is first presented → fall silent in the
presence of continued stimulation.

Rapid adapting afferents - effective in conveying info. about changes in

ongoing stimulation.

associated w. Pacinian corpuscles.

→ slowly adapting when corpuscles is removed.

Slowly adapting afferents - provide info. about the spatial attributes of the
stimulus.

Diff. in the properties of the channels - expressed in sensory afferents/ filter

properties of spec. receptor cells → encapsulate many sensory afferents.

generator potentials - produced only by restricted set of stimuli → impinge on a

given afferent fiber.

given sensory afferent - give rise to multiple peripheral branches → the transduction
properties of all branches of a single fiber are identical.

Somatosensory afferents - constitute parallel pathways → differ in conduction

velocity, RF size, dynamics & effective stimulus features.

Mechanoreceptors Specialised to receive Tactile Information

The distinct afferent pathway - to sensation → best developed for the glabrous
(hairless) portions of the hand.

Haptics -the interpretation of complex spatiotemporal patterns of stimuli → activate

many classes of mechanoreceptors.

Stereognosis - manipulate an ob. w. the hand → provide enough info. to identify the
ob.

recording the responses of indv. sensory afferents in human & non-human primate
→ poss. to characterise the responses of these afferents under controlled
conditions ~ gain insights into their contribution to somatic sensation.

Imp. aspect of the neurological assessment - invol. testing the functions of these
diff. classes of mechanoreceptive afferents

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 dermatomes - noting constrained zones → may present sensory loss in
patients w. nerve/ spinal cord injury.

Merkel cell afferents - slowly adapting fibers - account for about 25% of the
mechanosensory afferents in the hand.

→ enriched in fingertips ~ only that sample info from receptor cells.

Merkel cell-neurite complexes - in the tips of the prim. epidermal ridges -

extensions of the epidermis into the underlying dermis

→ coincide w. the prominent ridges ‘fingerprints’ on the finger surface.

Merkel cells - signal the static aspect of a touch stimulus.

play an active role in modulating the act. of their afferent axons ~ by releasing
neuropeptides on the neurites at junctions
→ resemble synapses w. the exocytosis of electron - dense secretory granules.

Merkel afferents - transduce the dynamic aspects of stimuli.

⇒ have highest spatial resolution of all sensory afferents - highly sensitive to

points, edges, and curvature → make them suited for processing info. about
shape & texture.

⇒ both Merkel cells + innervating sensory afferents - express the

mechanotransduction channel Piezo2 → sense stimuli.

delete Piezo2 in Merkel cells - reduces the sustained & static firing of the
innervating afferents.

Meissner afferents - rapidly adapting fibers - innervate the skin even more densely
than Merkel afferents → 40% of the mechanosensory innervation of the human
hand.

lie in the tips of the dermal papillae adjacent → prim. ridges and closest to the
skin surface.

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 → elongated receptors - formed by connective tissue capsule ~ contains a set of
flattened lamellar cells derived from Schwann cells + nerve terminals.

center of capsule - contains 2-6 afferent nerve fibers → terminate btwn &
around the lamellar cells.

configuration thought → contribute to the transient response of these

afferents to somatic stimulation.

Indentation of the skin - dynamic tension transduced by the collagen fibers -

provides the transient mechanical force

→ deforms the corpuscles ~ triggers generator potentials - induce a volley of

action potentials in the afferent fibers.

stimulus removed - indented skin relaxes → corpuscle return to its resting

configuration ~ generate other burst of action potentials.

⇒ Meissner afferents - display characteristic rapidly adapting (on-off

responses).

→ 4 times more sensitive to skin deformation as Merkel ~ larger RF than Merkel ⇒

transmit signal w. reduced spatial resolution.

effective in transducing info. w. low-freq. vibration - occur when ob. moved

across the skin.

💡 Info. conveyed by Meissner afferents - responsible for detecting slippage

btwn the skin + ob. field in hand
→ essential feedback info. for the efficient control of grip.

Pacinian afferents - rapidly adapting fibers → make up 10-15% of the

mechanosensory innervation in the hand.

located deep in the dermis/ subcutaneous tissue.

laminar capsule - concentric layers of membranes surrounding a single afferent

→ act as filter - allow only transient disturbances at high-freq. ~ activate the
nerve endings.

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 → Pacinian corpuscles - adapt more rapidly than Meissner ~ have a lower response
threshold.

(so sensitive - large RF - hard to define their boundaries)

⇒ detect vibrations transmitted thru. ob. - contact the hand/ being grasped in the
hand ~ esp. making/ breaking contact.

(imp. for skilled use of tools)

Ruffini afferents - slowly adapting fibers - least understood of the cutaneous

mechanoreceptors.

located deep in the skin ~ as well as ligament & tensions.

sensitive to the cutaneous stretching produced by digit/ limb movements →

20% of the mechanoreceptors in the human hand.
(responsive to skin stretches - during movement of fingers)

→ info. provide an accurate representation of finger position and the conformation

of the hand.

The microgeometries of different fine textures produce different

patterns of vibrations on the skin while the finger is scanning across
the textured surface, which are best detected by the rapidly
adapting afferents.

The longitudinal lanceolate endings - derived from Ab, As, C filters → form rapidly
adapting low-threshold mechanoreceptors associated w. the hairs.
→ imp. for mediating forms of sensual touch.

these responses - distinguised from responses of free nerve endings in the

epidermis ~ also derived from As & C axons in peripheral nerves.

these free nerve endings and the distinct fibers from which they are
derived have different physio- logical properties and respond (to

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 painful stimuli) at much higher activation thresholds than touch-
sensitive receptors associated with hair follicles.

Mechanoreceptors specialised for proprioception

Cutaneous mechanoreceptors - provide info. derived from external stimuli

another major class - provides info. about mechanical forces arising w/in the
body itself ~ part. from the musculoskeletal system.

The purpose of these proprioceptors → give detailed & continuous info. about the
position of the limbs + other body parts.

Low-threshold mechanoreceptors → provide this kind of sensory info. ~

essential to the accurate performance of complex movements.

Specialised proprioceptors - exist in the heart + major blood vessels → provide

info. about blood pressure BUT these neuron - considered to be part of the
visceral motor system.

Muscle spindles - found in all but a few striated (skeletal) muscles.

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 8 spec. intrafusal muscle fibers - surrounded by a capsule of connective tissue.

→ distributed among + parallel arrangement w/ the extrafusal fibers of skeletal

muscle → the true force-producing fibers.

spec. to signal changes in muscle length ~ low-threshold mechanoreceptors in

tendons → inform the CNS about changes in muscle tension.

Golgi tendon organs - formed by branches of group Ib afferents

distributed among the collagen fibers → form the tendons - provides
accurate samp. of the tension ~ exist in the muscle.

Sensory afferents - coiled around the central part of the intrafusal spindle
~ when muscle stretched → tension on intrafusal fibers - activates gated ion
channels in the nerve endings ~ triggering action potentials.

⇒ Innervation of muscle spindle - arise from 2 classes of fibers (prim. and secondary
endings).

1. Prim. endings - arise from the largest myelinated sensory axons (Ia afferents) →
rapid adapting responses to change in muscle lengths.
⇒ transmit info. about limb dynamics - the velocity and direction of movement.
2. 2nd endings (II afferents) - produce sustained responses to constant muscle
lengths.

⇒ provide info. about the static position of limbs.

The intrafusal fibers - contractile muscle fibers ~ controlled by a separate set of

motor neurons (y motor neurons) in the ventral horn of the spinal cord.

Changes in the tension of intrafusal fibers - sig. impact on the sensitivity of the
spindle afferents → changes in muscle length.

⇒ the level of act. in the y system must be taken - for central circuits to provide
accurate accounts of limb position & movement.

Large muscles - generate coarse movements have relatively few spindles.

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 Extraocular muscles + intrinsic muscles of the hand and neck - richly
supplied w. spindles
→ reflect the imp. of accurate eye movements + the need to manipulate ob. w. great
fitnesse + continuous demand for precise positioning of the head.

The rela. btwn. receptor density & muscle size - consistent w. the generalisation

→ sensorimotor apparatus at all levels of the nervous system is much richer (for
hands, head, speech organs, other parts of body).

💡 exp. w. vibrators - stimulate the spindles of spec. muscles → provided

compelling evidence - act. of afferents give rise to vivid sensations of
movement in immobilised limbs.

→ indicate that somatic and visual cues also play imp. roles.

Joint receptors - resemble many of the receptors foundin skin (incl. Ruffini endings
& Pacinian corpuscles). ⇒ imp. for judging position of the fingers.
→ playing protective role in signalling positions - lie near the limits of normal finger
joint range of motion.

Ruffini afferents + input from muscle spindles - contribute to fine representation

of finger position.

Central Pathways Conveying Tactile Information from the Body:

The Dorsal Column–Medial Lemniscal System

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 Both branches give off axonal collaterals - project into the gray matter of the spinal
cord across several adjacent segments → terminating in the deeper layers (laminae
III, IV, V) in the dorsal horn.

Dorsal columns (posterior funiculi) - main ascending branches extend

ipsilaterally → synapse on neurons in the dorsal column nuclei.

column - the columnar appearance of these fibers → run the length of the
spinal cord.

first-order neurons in the pathway - have quite long axonal processes →

neurons innervating the lower extremities ~ axons extend from their peripheral
targets thru. much of the length of the cord to the caudal brainstem.

Projection neurons located in laminae III, IV and V of the dorsal horn

→ receive inputs from mechanosensory collaterals project in parallel thru.
the dorsal column → to the same dorsal column nuclei.
(indirect mechanosensory → cross the midline - form a dorsoventrally
elongated tract (medial lemniscus))

The medial lemniscal axons - carry info. from the lower limbs located
ventrally/ axons related to upper limbs located dorsally.

axons synapse w. thalamus neurons - located in ventral posterior lateral

nucleus (VPL) → receive input from contralateral dorsal column nuclei.

Third-order neurons in VPL - send axons via internal capsule → terminate in

ipsilateral postcentral gyrus of cerebral cortex (prim. somatosensory cortex).
~ also send axons to 2nd somatosensory cortex.

💡 Somatosensory cortex - represents mechanosensory signals → generate in

the cutaneous surfaces of the contralateral body.

The Trigeminothalamic System

Cutaneous mechanoreceptor info. from the face → conveyed centrally by a

separate set of first-order neurons - located in trigeminal (cranial nerve V)

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 ganglion.

ophthalmic

maxillary

mandibular

⇒ subdivisions of the trigeminal ~ ea. innervates a well-defined territory on the face

and haed incl. teeth & mucosa of the oral & nasal cavities.

Central ganglion → form the sensory roots of the trigeminal nerve - enter the
brainstem at the level of the pons → to terminate on the neurons in the trigeminal
brainstem complex.

Most of the afferents → convey info. from low-threshold cutaneous

mechanoreceptors - terminate in principal nucleus.
→ nucleus corresponds to the dorsal column nuclei - relay mechanosensory
info. from the rest of the body.

Spinal nucleus - contains several subnuclei → receive inputs from collaterals of

mechanoreceptors.
→ trigeminal neurons - sensitive to pain, temperature, non-discriminative touch
project to the spinal nucleus.

Central Pathways Conveying Proprioceptive Information

from the Body

The axons of proprioceptive afferents - enter the spinal cord thru. the dorsal roots ~
many fibers from proprioceptive afferents
→ bifurcate into ascend + descending branches ~ send collateral branches to
several spinal segments.

The ascending branches of proprioceptive axons → travel w. the axons convey

cutaneous mechanosensory info. thru. the dorsal column.

info. supplied by proprioceptive afferents - imp. for ability to sense limb position ~
essential for the functions of the cerebellum

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 → regulates the timing of muscle contractions necessary for the performance of
voluntary movements.

Association w. cerebellar pathway → conveys proprioceptive info. for the

lower part of the body to dorsal column nuclei.

First-order proprioceptive afferents - enter spinal cord btwn mid-lumbar &

thoracic level - travel via dorsal column up to medulla → synapse on neurons in
Clarke’s nucleus (located in the medial aspect of the dorsal horn).

Second-order neurons in Clarke’s nucleus - send their axons into the ipsilateral
posterior lateral column of the spinal cord → travel up to the level of the medulla
in the dorsal spinocerebellar tract.

Third-order neurons - decussate + join in the medial lemniscus → accompany

the fibers from cutaneous mechanoreceptors in their course to VPL of the
thalamus.

from the Face

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 The cell bodies of the first order proprioceptive neurons for the face → found w/in
CNS in the mesencephalic trigeminal nucleus.

These pseudounipolar neurons - have peripheral processes → innervate muscle

spindles & Golgi tendon organs associated w. facial musculature + central
processes
→ project to brainstem nuclei responsible for reflex control of facial muscles.

Somatosensory components of the thalamus

Ea. of the serval ascending somatosensory pathways → converges on the ventral

posterior complex of the thalamus ~ terminates in an organised fashion.

VPL receives projections from the medial lemniscus - carrying somatosensory

info. from the body & posterior head.

Ventral posterior medial nucleus (VPM) - receives axons from the trigeminal
lemniscus → convey somatosensory info. from the face.

⇒ Info. supplied by diff. somatosensory receptors - remains segregated in its passage

to cortical circuits.

Primary somatosensory cortex

The major. of axons arise from neurons in ventral posterior complex of the thalamus
→ project to cortical neurons in layer 4 of prim. somatosensory cortex.

The prim. somatosensory cortex in humans - located in the postcentral gyrus of

the parietal lobe ~ comprise 4 distinct regions (Brodmann’s areas 3a, 3b, 1,2)

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 💡 Exp. of rat brain

inordinate amount of the somatosensory cortex - devoted to representing

the large facial whiskers ~ key components of the somatosensory
input for rats & mice.

⇒ Sensory input - sig. to given species gets relatively more cortical

representation.

These diff. in responses prop. - reflect parallel sets of inputs. from functionally
distinct classes of neurons in the ventral posterior complex.

A rich pattern of corticocortical connections btwn SI areas → contributes sig. to the

elaboration of SI response properties.

This arrangement of connections - establishes a functional hierarchy → 3b serves

as an obligatory first step in cortical processing of somatosensory info.

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 lesions of area 3b in non-human primates result in profound deficits
in all forms of tactile sensations mediated by cutaneous mecha-
noreceptors, while

lesions limited to areas 1 or 2 result in partial deficits and an

inability to use tactile information to discriminate either the texture
of objects (area 1 deficit) or the size and shape of objects (area 2
deficit).

VM - studies of finely spaced electrode penetration in 3b → provided strong

evidence - demonstrate neurons w. rapidly ~ slowly adapting prop. were clustered
into separate zones w/in the representation of a single digit.

the cortical slowly adapting neurons - show large touch OFF response in
addition to sustained firing during contact.
→ OFF response signalled only by rapidly adapting afferents in fingers.

⇒ The cortical rapidly + slowly dapting columns - reflect diff. processing of

convergent inputs from diff. peripheral receptors → convey distinct physiological
signals.

Beyond SI: Corticocortical and Descending Pathways

Somatosensory info. → distributed from the prim. somatosensory cortex to ‘high-

order’ cortical fields.

SII (2nd somatosensory cortex) - receives convergent projections from all

subvisions of SI ~ necessary for SII function.

→ IF lesions of SI → eliminate the responses of SII neurons ⇒

SII sends
projections in turn to limbic structures (amygdala & hippocampus).

SI also project to parietal areas posterior to area2, esp. 5a & 7b.

→ receive direct projections from area2 - supply inputs to neurons in motor &
premotor areas of the frontal lobe.

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 The projections from parietal cortex → motor cortex - critical for the integration of
sensory & motor info.

⇒ These pathways originate in sensory cortical fields → run to the thalamus, brainstem,
and spinal cord.
⇒ Descending projections from somatosensory cortex - outnumber ascending
somatosensory pathways → modulate the ascending flow of sensory info. at the level of
the thalamus & brainstem.

Plasticity in the adult cerebral cortex

K&M - examine the impact of peripheral lesions on the topographic maps in

somatosensory cortex.
→ corresponding region of cortex - found to be unresponsive ~ after few weeks -
became responsive to stimulation of neighbouring regions of the skin.

“functional remapping” - occurs in the somatosensory nuclei in the thalamus &

brainstem.
→ some reorganisation of cortical circuits - depend on this concurrent subcortical
plasticity.

(adjustment may contribute to the altered sensation of phantom limbs after

amputation)

⇒ reorganise after peripheral deprivation/ injury - general property of the mature

neocortex.

Appreciable changes in cortical representation - occur in response to physiological

changes in sensory/ motor exp.

⇒ Sig. changes in RF of somatosensory neurons - can be detected when peripheral

nerve is blocked by a local anesthetic.

The transient loss of sensory input from small skin area → induces a reversible
reorganisation of the RF of both cortical & subcortical neurons.
→ neurons assume new RF - respond to tactile stimulation of the skin surrounding
the anaesthetised region.

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 ⇒ Changes may be of limited value for recovery of function following brain injury - lead
to symptoms that detract ~ rather than enhance the quality of life following damage.

changes in cortical function - reflect alterations in the strength of synapses already

present
→ prevent/ redirect the synaptic events - underlie injury-induced plasticity could
reduce the long-term impact of acute brain damage.

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