Somatosensory System I: Rapidly adapting (RA) receptors

Tactile Discrimination and Position Sense o Transient, phasic, or vibratory stimuli

o Respond to initial application or removal of
Somatosensory System stimulus, but not to maintained stimulation
Transmits and analyzes touch or tactile information o Meissner, some hair follicle receptors, Pacinian
Modalities: Slowly adapting (SA) receptors
o Discriminative touch o Tonic events
o Flutter-vibration o Displacement, velocity, intensity, duration
o Proprioreception (position sense) o Active as long as the stimulus is present
o Crude (nondiscriminative) touch o Merkel, Ruffini, some hair follicle receptors
o Thermal (hot and cold) sensation o Merkel cell complexes for reading Braille
o Nociception (pain) Receptor (RA/SA) Sensation Receptive No. per cm
2

Pathways: Field Size FT Palm

o Posterior column-medial lemniscal pathway Meissner corpuscle Tap, flutter Small (54.9 >100 40
o Trigeminothalamic pathways 2
(RA) 5-40 Hz 8.6 mm )

Portions originating in the principal Hair follicle receptors Motion/ N/A N/A N/A
trigeminal sensory nucleus (RA/SA) direction
o Spinocerebellar pathways Pacinian corpuscle Vibration Large 20 10
(RA) 60-300 Hz
o Anterolateral system
Merkel cell (SA) Touch - Small 70 30
2
pressure (44.7 mm )
I. Posterior Column-Medial Lemniscal System (PCMLS) Ruffini complex (SA) Unknown Large 50 15
Perception/appreciation of mechanical stimuli Fiber Type: II FT - fingertip
o Fine form and texture discrimination
o Stereognosis (form recognition of 3D shape) Deep Tactile Mechanoreceptors
o Proprioception (conscious awareness of body Dermis, fascia surrounding muscles and bone,
position) peridontium, mesenteries
o Kinesthesia (limb movement in space) Pacinian corpuscles, Ruffini corpuscles, other encapsulated
General somatic afferent (GSA) fibers nerve endings
o Fast conduction velocity Pressure, vibration, skin stretch and distention, tooth
o Limited number of synaptic relays displacement
o Precise somatotopic organization
- Accurate localization Proprioceptive receptors
Limited convergence Muscles, tendons, joint capsules
o High fidelity Nuclear bag, nuclear chain intrafusal muscle fibers of
o High degree of spatial and temporal resolution muscle spindles and associated nerve fibers, Golgi tendon
Frequency coding organs and group 1b fibers, Ruffini-type joint receptors
o Firing rate signals stimulus intensity or temporal Static limb and joint position, dynamic limb movement
aspects (kinesthesia)
Population coding Balance, posture, limb movement
o Distribution in time and space of activated cells
in the CNS signals stimulus location, motion and Receptor (RA/SA) Sensation Function/ Signal FT
direction Nuclear bag fiber* High dynamic Length and rate Ia
1 annulospiral endings sensitivity of change; length
Lateral (surround) inhibition
and velocity
o Posterior column nuclei
Nuclear chain fiber* Low dynamic Length; tension II
o Present in all pathway relays 2 flower spray endings sensitivity
o Two-point discrimination Golgi tendon organ* Tension Muscle force; Ib

Sharpens and enhances the tendon
discrimination between separate Ruffini endings* Limb position Joint movement I
points on the skin and pressure
* Slowly adapting FT = Fiber Type
Peripheral Mechanoreceptors
Primary Afferent Fibers
Mechanical pressure  transduction (peripheral process of 1
afferent neuron  graded membrane potential  depolarizes Consist of:
trigger zone (at axon hillock) to threshold  action potential (1) Peripheral process from posterior root ganglion
* Merkel cells no transduction (affects 1 aff. neuron via NT) (PRG) to peripheral mechanoreceptors or free
nerve endings
Cutaneous Tactile Receptors (2) Central process from PRG to CNS
(3) Pseudounipolar cell body in the PRG
Basal epidermis, dermis of glabrous and hair skin
Peripheral distribution delineates segmental pattern of
Low-threshold
dermatomes (1 association: pain and temperature)
Encapsulated: Meissner, Pacinian, Ruffini corpuscles
Classification:
Unencapsulated: Merkel cell neurite complexes; hair
(1) Contribution to compound AP (A, B, C waves)
follicle receptors
(2) Specific to cutaneous fibers:
Greatest density: glabrous digits, perioral region
Based on fiber diameter, myelin thickness, and
Receptive field area of skin innervated by branches of a
conduction velocity (I, II, III, IV)
GSA fiber, the stimulation of which activates its receptors
Conduction velocity determines contribution to cmpd. AP
o Inverse rel. between size of receptive field and
Group 1a, 1b, II: discriminative touch, vibratory and
representation in the somatosensory cortex
position sense
Spinal Cord and Brainstem
Large-diameter (groups 1a, 1b, II, and A)
o Discriminative touch
o Flutter-vibration
o Proprioception
Enter the spinal cord through the medial division of the
nd
posterior root, branch into (1) 2 order neurons in the SC
grey matter  spinal reflexes; ascending projections (e.g.,
postsynaptic posterior column fibers; (2) gracile and
cuneate fasciculi (posterior columns) largest set
Postsynaptic Posterior Column Pathway
Nondiscriminative tactile signals to supraspinal levels
Non-primary afferent axons carrying tactile signals in PC
Origin: Laminae III, IV of posterior horn
Axons of second-order postsynaptic PC fibers travel in the
PC columns and terminate in the PC nuclei
o Relayed to contralateral thalamus via the
medial lemniscus
Possible morphological basis for return of some tactile
sensation after PCMLS vascular lesions

Posterior columns Classification Diameter Cond. Receptor Supplied

Topographic organization Contribution Afferent (m) Velocity
Sacral level fibers medial to cmpd. AP fibers only (m/s)
o Fibers up to T6 are added laterally to form Sensory Fiber Type
gracile fasciculus (medial) A Ia and Ib 13-20 80-120 Primary muscle
o Cervical fibers, T1-T5  cuneate fasciculus spindles, Golgi
(lateral) tendon organ
o Lower extremity is represented medially and vice A II 6-12 35-75 Secondary muscle
versa spindles, skin
Compromised blood flow in the posterior spinal artery mechanoreceptors
(supplies posterior funiculus), or mechanical injury to A III 1-5 5-30 Skin
posterior columns (e.g., Brown-Sequard syndrome)  mechanoreceptors,
ipsilateral reduction/loss of TdVP sensations at and thermal receptors,
below segmental level of injury and nociceptors
o Also seen in tabes dorsalis (progressive C IV 0.2-1.5 0.5-2 Skin
locomotor ataxia) d/t Treponema pallidum mechanoreceptors,

Associated with neurosyphilis thermal receptors,
and nociceptors
Posterior column nuclei (gracile, cuneate nuclei) Motor Fiber Type
Posterior medulla, at end of respective fasciculi A N/A 12-20 72-120 Extrafusal skeletal
Supplied by posterior spinal artery muscle fibers
Cell bodies are second-order neurons in the PCMLS A N/A 2-8 12-48 Intrafusal muscle
st fibers
Receive input from 1 -order neurons with cell bodies in
the ipsilateral PRG B N/A 1-3 6-18 Preganglionic
Rapidly adapting inputs terminate centrally and caudally autonomic fibers
Slowly adapting inputs cutaneous, muscle spindle and C N/A 0.2-2 0.5-2 Postganglionic
joint; project to rostral pole of PC nuclei and to rostrally autonomic fibers
adjacent nucleus z
Also receive descending axons from contralateral 1 Ventral Posterolateral Nuclei
somatosensory cortex and medullary reticular formation VPN (ventrobasal complex) wedge-shaped cell group
(nucleus reticularis gigantocellularis) located caudally in the thalamus
Inner core large projection neurons Composed of:
Shell small fusiform and radiating cells; interneurons (1) Ventral posterolateral nucleus (VPL)/
responsible for feedback inhibition (alters activity of Ventralis caudalis externus
projection neurons of the inner core) o Ascending input from medial lemniscus
Info received by PC nuclei undergoes signal processing (2) Ventral posteromedial nucleus (VPM)/

nd
2 order cells in the inner core send axons to Ventralis caudalis internus
contralateral thalamus o Input from trigeminothalamic tracts
Internal arcuate fibers * Separated by arcuate lamina
o Located in the medulla Supplied by thalamogeniculate branches of the posterior
o Axons of PC nuclei cells cerebral artery (compromise can result in loss of all tactile
o Arc anteromedially toward midline, decussate, sensation over contralateral body and head)
and ascend as contralateral medial lemniscus

Cuneate posterior Ventral Posterolateral Nucleus (VPL)

Gracile anterior Somatotopic organization: Lower extremity lateral
Upper extremity medial
Medial Lemniscus Representation of individual body part: C-shaped lamina
Supplied by anterior spinal artery in the medulla; Tactile signals also represented in other thalamic nuclei
penetrating branches of the basilar artery in the pons with lemniscal input (e.g. ventral posterior inferior (VPI) n.;
Vascular damage leads to contralateral TdVP deficits pulvinar and lateral posterior groups)
Rotates as it moves laterally such that: Functional organization:
o Upper extremity is represented medially Core: RA and SA inputs
o Lower extremity is represented laterally Shell: Pacinian inputs; inputs from joints and muscles
Terminates in ventral posterolateral nucleus (VPL) of the * posterior, rostral, and anterior edges of n.
thalamus Rods: sagittal longitudinal cell clusters where axons arborize
o Isorepresentation arranged along rostracaudal axis
 Cell populations identified in the VPL for the trunk and
extremities, VPM for the head:
(1) Thalamocortical cells and fibers
o Third-order neurons
o Excitatory (glutaminergic) input to the cortex
o Large-diameter multipolar cells; axons traverse
the posterior limb of the internal capsule
o Terminate in the primary (SI) and secondary (SII)
somatosensory cortices
(2) Local circuit interneurons
o Inhibitory (GABAergic)
o Receive excitatory corticothalamic inputs and
influence the firing rate of thalamocortical cells
o Influenced by:
GABAergic input from the thalamic reticular n.
Excitatory (glutaminergic) corticothalamic fibers
fr. Layer VI of 1 and 2 somatosensory cortices
Primary Somatosensory Cortex (SI)
Bordered anteriorly by central sulcus, posteriorly by
postcentral sulcus; comprises postcentral gyrus and
posterior paracentral gyrus
Contains somatotopic representation of the body surface
Cortical representation is directly proportional to receptor
density
Vascular lesions involving the:
Middle cerebral a. Contralateral upper body and face
Anterior cerebral a. Contralateral lower limb
Histologic Division
BA 3a
o Depths of the central sulcus
o Abuts BA 4 (1 motor cortex)
BA 3b, 1
o Bank of sulcus to shoulder of postcentral gyrus
BA 2
o Gyral surface
o Abuts BA 5 (somatosensory association cortex)
BA 3a, 2
o Shell neurons
o Proprioceptive inputs from muscle spindle
afferents (BA 3a), Golgi tendon organs, and joint
afferents (BA 2)
o Kinesthetic information related to muscle length
and tension; static and transient joint position
BA 3b, 1
o Core neurons
o Cutaneous inputs from Meissner corpuscles (RA)
and Merkel cells (SA), and pain/thermoreceptors
Lesion Deficit
BA 1 Texture discrimination
BA 2 Size and shape discrimination (astereognosis)
BA 3b Texture, size and shape discrimination*
Lesions frequently include larger areas, resulting in more global
deficits (e.g., proprioception, position sense, vibratory sense,
and pain and thermal sensations on the contralateral side)
* Suggest hierarchical info processing (3b performs 1 processing)
Additional Cortical Somatosensory Regions
Secondary Somatosensory Cortex (SII)
Inner face of the upper bank of the lateral sulcus
Somatotopic representation
Inputs from ipsilateral SI cortex, VPI n. of thalamus
(anterior/ventral to VPL/VPM)
Supplied by middle cerebral artery
BA 5, Lateral Portions of BA 7 (7b)
Inputs from anterior and lateral posterior group
o Inputs from medial lemniscus and SI
Parietal Association Area Lesions
Agnosia contralateral body parts are lost from the
personal body map
o limb is not dressed or recognized as part of
patients own body
Trigeminal System
Info from craniofacial structures (inc. oral, nasal cavities)
Primary afferent neurons of trigeminal nerve  brainstem
nuclei (in column which begins rostrally in the middle pons
and ends to overlap with the posterior horn of the upper
cervical spinal cord)  pathways
Tactile discrimination, flutter-vibration, proprioception,
pain and thermal sensations
Trigeminal Nerve
(1) Ophthalmic (V1)
(2) Maxillary (V2)
(3) Mandibular (V3)
Also receives GSA input from CN VII, IX, X small
cutaneous area around the ear
Cell bodies of 1 afferent neurons are located in the
trigeminal (gasserian or semilunar) ganglion and
mesencephalic trigeminal nucleus
Central processes form sensory root (portio major) and
enter lateral aspect of pons
o Bifurcate into ascending and descending branches
before terminating on 2-order neurons in the
brainstem trigeminal sensory nuclei

Ascending  principal sensory n. (pons)

Descending  spinal (descending) tract
of the trigeminal nerve  spinal
nucleus of the trigeminal nerve
II. Anterior and Posterior Trigeminothalamic Tracts
Peripheral Receptors
Similar to those found in other parts of the body
May serve specialized functions
o Receptors in the periodontal ligament:
tooth displacement and bite force
o Encapsulated receptors (primarily Meissner)

Lips, perioral skin
Axons are mostly large-diameter
May ascend without branching, or bifurcate before entering
the principal sensory nucleus
Principal sensory nucleus
o In the middle pons, at rostral pole of spinal
trigeminal nucleus
o Dorsomedial division

Input from oral cavity
o Ventrolateral division

Trigeminal nerve (all components)

Inverted when viewed in the anatomic
orientation; upright when viewed in the
clinical orientation (e.g., MRI or CT):
V1 anterior
V2 middle
V3 posterior
- Second-order neurons

Relatively small receptor fields

Subject to same type of intranuclear
modulations as PC nuclei cells
 Ventrolateral neuron axons + fibers from the spinal
trigeminal nucleus  trigeminal lemniscus/ anterior
(ventral) trigeminothalamic tract  contralateral VPM
o Courses near the medial lemniscus
Dorsomedial neurons  posterior (dorsal)
trigeminothalamic tract)  ipsilateral VPM
o Ascends in the pontine tegmentum lateral to the
periaqueductal gray, in association with the
central tegmental tract
Somatotopic representation in the VPM
o Oral cavity  medial
o External facial structures  lateral
Third-order thalamocortical neurons in the VPM 
posterior limb of internal capsule 
laterally-placed face area of SI in the postcentral gyrus
Perioral regions highest peripheral innervation density
 largest representation
Trigeminal mesencephalic nucleus
o Primary afferent neurons innervate muscle
spindles (proprioceptive) in jaw muscles and
some modified Ruffini endings and free nerve
endings in periodontal ligaments
o Slender column of pseudounipolar cells
o Inputs from peptidergic and monoaminergic
neurons in the brainstem

Presynaptic modulation before central
relay of primary afferent information
o Cell processes form the mesencephalic tract of
the trigeminal nerve

Adjacent to mesencephalic nucleus;
extends rostrally to border the
midbrain aqueductal gray

Central processes innervate trigeminal
motor nucleus cells  afferent limb of
the myotatic jaw jerk reflex
Myostatic Jaw Jerk Reflex
Processes of trigeminal mesencephalic n. neurons
Innervate muscle spindles in jaw-closing muscles
(monosynapc)
Trigeminal motor neurons
Innervates temporalis (elevates the jaw)
Gentle tap on the jaw activates afferent fibers  contraction of
homonymous muscle (temporalis) and synergists (masseter)
Periodontal ligament afferents feedback to jaw muscle
motor neurons during mastication to regulate bite force
o NOT monosynaptic
Probst tract
o Descending branches of mesencephalic tract fibers
o Distributes to the reticular formation, spinal
trigeminal nucleus, and cerebellum
Coordination of oral motility patterns (mastication,
swallowing, speech); oral stereognosis
Provides proprioceptive input to the principal sensory
nucleus and spinal trigeminal nucleus (d/t bifurcation)
o Receptors may be innervated by trigeminal
ganglion cells (e.g., temporomandibular joint,
extraocular muscles, periodontal ligaments)
o Principal sensory nucleus  cerebellum, spinal
cord, thalamus

Disproportionate share of large-
diameter, heavily myelinated fibers

Trigeminal homologue of the PC nuclei
Receptive Field Properties of Cortical Neurons
Cortical columns
Functional units into which neurons in cortical areas
representing the body and head are organized
Distributed from the pial surface to the white matter
Cells in a column have similar peripheral receptive field loci
Thalamocortical inputs  layer IV, lower parts of layer III of
the SI cortex  pyramidal cells within individual columns
Cortical neurons
Complexity: cortical > subcortical neurons
Respond to a specific stimulus orientation (edges) and
specific textures
Can code for velocity, speed, and direction of moving stimuli
Three populations which receive proprioceptive inputs
(1) Simple: single joint or muscle group

RA, signal movement
(2) Postural

Signal final joint position when
movement is completed
(3) Multijoint: several joints and muscle groups

Signal complex multiple-joint
interactions
Larger receptive fields; more complex inhibitory surrounds
than displayed by subcortical inputs
Sensory signal processing can include:
(1) Convergence of afferent input
- Increases receptive field size, decreases resolution
(2) Divergence of output signal
- Relay cells can amplify the sensory signal and
apply it to multiple targets
(3) Facilitation
(4) Inhibition
** Enhance the signal-noise ratio in terms of both space and time
Neuroimaging and Functional Localization
Functional magnetic resonance imaging (fMRI)
o BA 3b, 1 increased blood flow in response to
air puff stimulation on the upper limb
Regional cerebral blood flow (rCBF)
Positron emission tomography (PET)
o BA 3b, 1 discrimination of moving stimuli
o BA 2 palpating shape and curvature
o Tactile stimulus activates SI and SII cortices
Magnetoencephalography (MEG)
Plasticity and Reorganization in the Primary Somatosensory Cortex
Brain development
Child
Malleable/plastic
Pruning occurs during a critical period
o Many will be retained
o Some will be pruned through apoptosis
Plasticity
o Ability to reassign brain functions to other brain
regions
Adult
Relatively non-malleable
Somatosensory cortex can undergo reorganization
o Changes in cortical map after amputation:
expansion of cortical representation to adjacent
parts (begins within 10 days after amputation)

phantom limb
Stroke better prognosis for younger patients
o The older the brain, the less apparent plasticity
III. Nonconscious Proprioception: Spinocerebellar Pathways Trigeminal Connections
Information about limb position, joint angles, muscle Proprioceptive information from jaw muscle spindles,
tension and length periodontal afferents, and the temporomandibular joint
o Cerebellar control of body muscle tone, o Involved in mastication, long-range functional
movement, and posture Spinocerebellum changes (e.g., transition from suckling to
Spinocerebellar  cerebellar nuclei chewing in the newborn; natural dentition to
tract axons  mossy fibers  vermis and paravermis denture use)
o Somatotopic representations in the anterior lobe Branches of the mesencephalic
and paravermis of posterior lobe trigeminal neurons  superior head
Degeneration: Friedreich ataxia  cerebellar ataxia cerebellar peduncle  cerebellum representation
o Lack of coordination d/t lack of sensory feedback Spinal trigeminal nucleus
pars interpolaris, pars caudalis 
Posterior Spinocerebellar Tract (PSCT) restiform body  cerebellum
Proprioreceptive + some exteroreceptive (cutaneous) afferents in the
lower limb and lower trunk**
Posterior root fibers
Dorsal nucleus of Clarke*
(In lamina VII of the intermediate zone of spinal segments T1-L2)
Ipsilateral lateral funiculus
Surface of the SC, lateral to corticospinal tract
Restiform body
Ipsilateral cerebellar cortex

* Primary afferent fibers from spinal levels caudal to L2 ascend in

the posterior funiculus to reach the DNC
** Group I muscle spindle, Golgi tendon organ
Monosynaptic activation
Discharge rate has a linear relationship with muscle length
(Firing rate encodes muscle length as a frequency code)
** Group II and III tactile fibers

Cuneocerebellar Tract
(Upper limb equivalent of PSCT)
Muscle spindle and exteroreceptive afferents**
Posterior root fibers in spinal segments C2 to T4
Ipsilateral cuneate fasciculus
Lateral cuneate nucleus*
Cuneocerebellar fibers
Restiform body
Ipsilateral cerebellar cortex

* Also receives inputs from proprioceptive primary afferent fibers

** Exteroceptive input terminates in the folia of the anterior lobe in
lobule V

Anterior Spinocerebellar Tract (ASCT)

Group 1 afferents in the lower limb (L3-L5, in the lateral part of
Rexed laminae V-VII and anterolateral border of the anterior horn
[spinal border cells])
Cross the midline in the anterior white commissure
Lateral funiculus (anterior to the PSCT)
Pons turn posterolateral, superior cererebellar peduncle
Recross to terminate in the ipsilateral cerebellum
- More lateral distribution compared to PSCT

* Cells giving rise to ASCT fibers are influenced by

descending projections of the reticulospinal (inhibits) and
corticospinal (facilitates) pathways
** Vestibulospinal and rubrospinal projections  monosynaptically
excitates ASCT cells

Rostral Spinocerebellar Tract

(Upper limb equivalent of the ASCT)
Cutaneous tactile information from Meissner, Merkel and pacinian
mechanoreceptors (group II and group III afferents)
Cell bodies in lamina VII of the cervical enlargement (C4-C8)
Lateral funiculus of the spinal cord
Restiform body Superior cerebellar peduncle
Cerebellum