Sensory Pathways

SENSORY PATHWAYS
Sensory pathways include only those routes which conduct information to the conscious
cortex of the brain. However, we will use the term in its more loosely and commonly
applied context to include input from all receptors, whether their signals reach the
conscious level or not.

GENERAL SOMATIC AFFERENT (GSA) PATHWAYS FROM THE

BODY
Pain and Temperature
Pain and temperature information from general somatic receptors is conducted over
small-diameter (type A delta and type C) GSA fibers of the spinal nerves into the
posterior horn of the spinal cord gray matter (Fig-1). These are monopolar neurons with
cell bodies in the posterior root ganglia. After entering the cord, the fibers pass up or
down in the dorsolateral tract, located between the tip of the posterior horn and the
surface of the spinal cord near the posterior root, before finally synapsing in laminae III
and IV.

Fig-1

Sensory Pathways
Second-order neurons from these synapses cross over to the opposite side of the cord in
the anterior white commissure, where they turn upward as the lateral spinothalamic
tract (LSTT). At higher pontine levels this tract comes to lie close to the medial
lemniscus, with which it travels to the ventral posterior lateral nucleus (VPL) of the
thalamus. Some fibers of this tract don't enter the thalamus but end instead in the
brainstem reticular formation. After synapsing in the thalamus, third-order neurons enter
the posterior third of the internal capsule, pass through the corona radiata, and
terminate in the primary and secondary sensory areas of the parietal lobe cortex (areas
3,1, and 2). Notice that regardless of the level of entry into the spinal cord, pain and
temperature stimulation delivered to one side of the body registers in the cerebral cortex
of the opposite side.
Fast and Slow Pain
Pain sensation is often confusingly labeled "fast" or "slow" depending on the type of fiber
which conducts the impulse and the speed with which the signal consciously registers.
Fast pain, often called sharp or pricking pain, is usually conducted to the CNS over type
A delta fibers. These ultimately excite lateral spinothalamic tract fibers which go directly
to the VPL of the thalamus on the contralateral side. From here third-order fibers project
to the cerebral cortex where they are somatotopically organized and sharply localized.
Somatotopic organization means that each minute area of the sensory cortex receives
input from a distinct peripheral area. A person can sharply localize a pain if he is able to
tell exactly where it is originating. Slow pain, often called burning pain, is conducted to
the CNS over smaller-diameter type C fibers. After entering the cord these fibers
stimulate lateral spinothalamic tract neurons which send collaterals into the brainstem
reticular formation. Fibers from the reticular formation diffusely project to the thalamus,
hypothalamus, and possibly other areas as well, perhaps giving rise to the emotional
component of pain. Pain signals following this route are poorly localized.
Dermatomes
A dermatome is the area of skin supplied by the afferent fibers in the posterior root of a
single spinal nerve. Dermatomes tend to overlap each other so that stimulation of a
specific point on the skin typically sends afferent signals into the cord over more than
one posterior root. This is functionally important since destruction of a single posterior
root does not totally eliminate sensation from the afflicted dermatome.
Touch and Pressure
Touch can be subjectively described as discriminating or crude. Discriminating (epicritic)
touch implies an awareness of an object's shape, texture, three-dimensional qualities,
and other fine points. Also implied here is the ability to recognize familiar objects simply
by tactile manipulation. Crude (protopathic) touch, on the other hand, lacks the fine
discrimination described above and doesn't generally give enough information to the
brain to enable it to recognize a familiar object by touch alone. The tactile information
implied here is of a much cruder nature than described for epicritic touch. The pathways
to the brain for these two kinds of touch appear to be distinct.
Crude (Protopathic)
Touch and Pressure
General somatic mechanoreceptors sensitive to crude touch and pressure conduct
information into the cord over GSA nerve fibers (Fig-2). The fibers pass up or down a
few cord segments (neuromeres) in the dorsolateral (Lissauer) tract before synapsing
chiefly in laminae VI, VII, and VIII. Second-order neurons cross over to the opposite side

Sensory Pathways
in the anterior white commissure to the anterior funiculus, where they turn upward in
the anterior spinothalamic tract (ASTT) to the VPL of the thalamus. At higher pontine
levels the tract also comes to lie close to the medial lemniscus as it ascends to the
thalamus. Third-order neurons project from the VPL to areas 3, 1, and 2 of the cerebral
cortex. Some of the ASTT fibers send collaterals into the brainstem reticular formation.
While some of these no doubt ultimately reach the thalamus by reticulothalamic
projections, the principal fate and function of these collaterals is largely unknown.

Fig-2

Discriminating (Epicritic) Touch, Pressure, and Kinesthesia

The conscious awareness of body position and movement is called the kinesthetic sense.
It's important to recognize that there are many receptors throughout the body which
continually conduct information to the brain concerning the body's position and
movement and even the level of muscle tone. Such receptors are collectively called
proprioceptors. However, not all of these signals reach the conscious level as a large
portion are conducted instead to the brainstem and cerebellum for subconscious
evaluation and integration. Only those proprioceptive signals reaching the conscious
level contribute to the kinesthetic sense. The kinesthetic sense and discriminating touch
and pressure pathways share a common route to the brain (Fig-3).

Sensory Pathways

Fig-3

General somatic mechanoreceptors sensitive to discriminating touch and pressure and

body position and movement conduct signals into the cord over GSA fibers. They pass
directly into the ipsilateral posterior funiculus, where they turn upward in the dorsal
columns to terminate in the dorsal column nuclei of the medulla. Those fibers entering
the cord below the midthoracic level (i.e., from the lower trunk and legs) ascend through
the medial dorsal column as the fasciculus gracilis and terminate in the nucleus gracilis.
Fibers entering the cord above the midthoracic level (i.e., from the upper trunk and
arms) enter the more lateral dorsal column and ascend as the fasciculus cuneatus to
terminate in the more lateral dorsal column nuclei, the nucleus cuneatus. As might be
expected, the dorsal columns include the fasciculus gracilis and fasciculus cuneatus while
the dorsal column nuclei include the nucleus gracilis and nucleus cuneatus. Second-order
neurons from these nuclei cross over to the other side of the brainstem in the lower
medulla as the internal arcuate fibers. which then turn upward in the medial lemniscus
to the VPL of the thalamus. Third-order neurons then project through the posterior limb
of the internal capsule to areas 3, 1, and 2 of the cerebral cortex.
Much of the proprioceptive information which reaches the conscious level giving rise to
the kinesthetic sense originates in joint receptors. However, recent evidence indicates
that signals from muscle spindles may also represent a significant contribution to
kinesthetic sensation. On the other hand, the subconscious proprioceptive information

Sensory Pathways
which is shunted to the brainstem and cerebellum for evaluation and integration arises
chiefly in muscle spindles and Golgi tendon organs.
Subconscious Proprioception
Most of the subconscious proprioceptive input is shunted to the cerebellum. Further,
signals arising in proprioceptors on the left side of the body register on the left side of
the cerebellum. By contrast, sensory signals arising in the left side of the body register
on the right side of the cerebral cortex. After entering the cord, proprioceptive afferents
(GSA fibers) terminate in laminae V, VI, and VII (Clarke's column) of the posterior horn.
Second-order neurons (primarily conducting information from Golgi tendon organs) cross
over to the opposite side of the cord in the anterior white commissure to the lateral
funiculus, where they turn upward in the anterior spinocerebellar tract (ASCT). After
reaching upper pontine levels the fibers cross back over and enter the cerebellum
through the superior cerebellar peduncle, where they terminate in the vermis (Fig-4).
Some of the anterior spinocerebellar tract fibers upon reaching the medulla remain
uncrossed and enter the cerebellum via the inferior cerebellar peduncle and terminate in
the contralateral vermis. Other second-order neurons (those receiving information
primarily from muscle spindles and tendon organs) leave Clarke's column to ascend in
the ipsilateral posterior spinocerebellar tract (PSCT) to the cerebellum. After reaching
the medulla, the fibers enter the cerebellum via the inferior cerebellar peduncle to
terminate in the ipsilateral cortex.

Fig-4

Sensory Pathways
Some of the subconscious proprioceptive input from the cervical region follows an
alternate route to the cerebellum. Some of the fibers travel a short distance in the dorsal
funiculus, terminating in the accessory cuneate nucleus of the medulla. Second-order
neurons project from here as the cuneocerebellar tract to enter the cerebellum via the
inferior cerebellar peduncle.
Posterior Funiculus Injury
Certain clinical signs are associated with injury to the dorsal columns. As might be
expected, these are generally caused by impairment to the kinesthetic sense and
discriminating touch and pressure pathways. They include (1) the inability to recognize
limb position, (2) astereognosis, (3) loss of two-point discrimination, (4) loss of vibratory
sense, and (5) a positive Romberg sign. Astereognosis is the inability to recognize
familiar objects by touch alone. When asked to stand erect with feet together and eyes
closed, a person with dorsal column damage may sway and fall. This is a positive
Romberg sign.

GENERAL SOMATIC AFFERENT (GSA) PATHWAYS FROM THE

FACE
Pain, Temperature, and Crude Touch and Pressure
General somatic nociceptors, thermoreceptors, and mechanoreceptors sensitive to crude
touch and pressure from the face conduct signals to the brainstem over GSA fibers of
cranial nerves V, VII, IX, and X. The afferent fibers involved are processes of monopolar
neurons with cell bodies in the semilunar, geniculate, petrosal, and nodose ganglia,
respectively. The central processes of these neurons enter the spinal tract of V, where
they descend through the brainstem for a short distance before terminating in the spinal
nucleus of V. Second-order neurons then cross over the opposite side of the brainstem
at various levels to enter the ventral trigeminothalamic tract, where they ascend to the
VPM of the thalamus. Finally, third-order neurons project to the "face" area of the
cerebral cortex in areas 3, 1, and 2 (Fig-5).

Sensory Pathways

Fig-5

Discriminating Touch and Pressure

The pathway for discriminating touch from the face is illustrated in Fig-6. Signals are
conducted from general somatic mechanoreceptors over GSA fibers of the trigeminal
nerve into the principal sensory nucleus of V, located in the middle pons. Second-order
neurons then conduct the signals to the opposite side of the brainstem, where they
ascend in the medial lemniscus to the VPM of the thalamus. Thalamic neurons then
project to the "face" region of areas 3, I, and 2 of the cerebral cortex.

Sensory Pathways

Fig-6

Kinesthesia and Subconscious Proprioception

Proprioceptive input from the face is primarily conducted over GSA fibers of the
trigeminal nerve. Curiously, however, the cell bodies of these monopolar neurons are
located in the mesencephalic nucleus of V in the midbrain rather than the semilunar
ganglia, where the cell bodies of other afferent neurons of the trigeminal nerve are
located. The peripheral endings of these neurons are the general somatic
mechanoreceptors sensitive to both conscious (kinesthetic) and subconscious
proprioceptive input. Their central processes extend from the mesencephalic nucleus to
the principal sensory nucleus of V in the pons (Fig-7).

Fig-7

The subconscious component is conducted to the cerebellum, while the conscious

component travels to the cerebral cortex. Certain second-order neurons from the
principal sensory nucleus relay proprioceptive information concerning subconscious
evaluation and integration into the ipsilateral cerebellum. Other second-order neurons
project to the opposite side of the pons and ascend to the VPM of the thalamus as the

Sensory Pathways
dorsal trigeminothalamic tract. Thalamic projections terminate in the face area of the
cerebral cortex.

SPECIAL SOMATIC AFFERENT (SSA) PATHWAYS

Hearing
The organ of Corti with its sound-sensitive hair cells and basilar
membrane are important parts of the sound transducing system for
hearing. Mechanical vibrations of the basilar membrane generate
membrane potentials in the hair cells which produce impulse patterns
in the cochlear portion of the vestibulocochlear nerve (VIII). The
principles of this system will be examined elsewhere. For now we will
examine only the central pathways from the receptors to their
terminations in the brain (Fig-8).
Special somatic nerve fibers of cranial nerve VIII relay impulses from
the sound receptors (hair cells) in the cochlear nuclei of the brainstem.
These are bipolar neurons with cell bodies located in the spiral ganglia
of the cochlea. Their central processes terminate in the dorsal and
ventral cochlear nuclei on the ipsilateral side of the brain stem at the
pontomedullary border. Most of the second-order neurons arising in the
cochlear nuclei cross to the opposite side of the brainstem in the
trapezoid body and turn upward in the lateral lemniscus, terminating in
the inferior colliculus of the midbrain. Collaterals of the lateral
lemniscus terminate in the nucleus of the trapezoid body, superior
olivary nucleus, nucleus of the lateral lemniscus, and the brainstem

Fig-8

Sensory Pathways
reticular formation. Fibers arising in these nuclei also ascend in the
lateral lemniscus. Those fibers from the cochlear nuclei which don't
cross over in the trapezoid body ascend in the ipsilateral lateral
lemniscus to the inferior colliculus. Sound signals also pass from one
side to the other via contralateral projections from one lemniscal
nucleus to the other as well as from one inferior colliculus to the other.
Thus each lateral lemniscus conducts information from both sides,
which helps to explain why damage to a lateral lemniscus produces no
appreciable hearing loss other than problems with sound localization.
Signals are then conducted from the inferior colliculi to the medial
geniculate bodies and finally to the primary auditory area of the
temporal lobes (area 41).

Vestibular System
The vestibulocochlear nerve serves two quite different functions. The cochlear portion,
previously described, conducts sound information to the brain, while the vestibular
portion conducts proprioceptive information. It is the central neural pathways of the
latter function which we will examine now (Fig-9). The mechanics and physiology of the
system explained elsewhere.

Fig-9

Special somatic afferent fibers from the hair cells of the macula utriculi and macula
sacculi conduct information into the vestibular nuclei on the ipsilateral side of the pons
and medulla. These are bipolar neurons with cell bodies located in the vestibular
ganglion. Some of the fibers project directly into the ipsilateral cerebellum to terminate
in the uvula, flocculus, and nodulus, but most enter the vestibular nuclei and synapse
there.

Sensory Pathways
As might be expected, neuronal output from the vestibular nuclei effects bodily and eye
movements in response to movements of the head as detected by the vestibular
apparatus. The vestibulospinal path fibers which affect body reflexes and muscle tone in
response to vestibular input originate primarily in the lateral vestibular nucleus. The
medial vestibular nucleus is the principal origin of both crossed and uncrossed fibers
which descend through the brain stem in the medial longitudinal fasciculus to the upper
cord causing various reflex head and arm movements in response to vestibular stimuli.
Finally, all four vestibular nuclei (medial, lateral, superior, and inferior) project both
crossed and uncrossed fibers to the motor nuclei of cranial nerves Ill, IV, and VI in order
to control and coordinate reflex eye movements. These vestibuloocular paths also travel
in the medial longitudinal fasciculus.

Vision
The visual system receptors are the rods and cones of the retina. The neurophysiology of
vision and visual reflexes are discussed elsewhere. Special somatic afferent fibers of the
optic nerve (II) conduct visual signals into the brain. Examination of Fig-10 will show
that fibers from the lateral (temporal) retina of either eye terminate in the lateral
geniculate body on the same side of the brain as that eye. On the other hand, SSA II
fibers from the medial (nasal) retina of each eye cross over in the optic chiasm to
terminate in the contralateral lateral geniculate body. The optic nerve is composed of
fibers from the retina to the optic chiasm. Even though no synapses occur in the optic
chiasm, the continuation of the visual pathway from the optic chiasm to the lateral
geniculate body is called the optic tract rather than the optic nerve. After a synapse in
the lateral geniculate body, the signal continues in the optic radiation to area 17 of the
conscious visual cortex. Area 17 is the primary visual area, which receives initial visual
signals. Neurons from this area project into the adjacent occipital cortex (areas 18 and
19) which is known as the secondary visual area. It is here that the visual signal is fully
evaluated.

Fig-10

Fig-11

The visual reflex pathway involving the pupillary light reflex is illustrated in Fig-11. This
is the well-known reflex in which the pupils constrict when a light is shined into the eyes
and dilate when the light is removed. Some SSA II fibers leave the optic tract before
reaching the lateral geniculates, terminating in the superior colliculi instead. From here,
short neurons project to the EdingerWestphal nucleus (an accessory nucleus of III) in
the midbrain, which serves as the origin of the preganglionic parasympathetic fibers of
the oculomotor nerve (GVE III). The GVE III fibers in turn project to the ciliary ganglia,

Sensory Pathways
from which arise the postganglionic fibers to the sphincter muscles of the iris, which
constrict the pupils.

GENERAL VISCERAL AFFERENT (GVA) PATHWAYS

Pain and Pressure Sensation via the Spinal Cord
Visceral pain receptors are located in peritoneal surfaces, pleural membranes, the dura
mater, walls of arteries, and the walls of the GI tube. Nociceptors in the walls of the GI
tube are particularly sensitive to stretch and overdistension.
General visceral nociceptors conduct signals into the spinal cord over the monopolar
neurons of the posterior root ganglia. They terminate in laminae III and IV of the
posterior horn as do the pain and temperature pathways of the GSA system; however,
their peripheral processes reach the visceral receptors via the gray rami communicantes
and ganglia of the sympathetic chain (Fig12), Second-order neurons from the posterior
horn cross in the anterior white commissure and ascend to the thalamus in the anterior
and lateral spinothalamic tracts, Projections from the VPL of the thalamus relay signals
to the sensory cortex.

Fig-12

Fig-13

Sensory Pathways

The localization of visceral pain is relatively poor, making it difficult to tell the exact
source of the stimuli. At least a partial explanation of our inability to precisely localize
visceral pain relates to its rarity. True visceral pain seldom occurs when compared to the
frequency of external pain. An additional compounding factor is the phenomena of
referred pain. Because true visceral pain is often projected or "referred" by the brain to
some area on the surface of the body, its true visceral origin is often confused. The
mechanism for referred visceral pain is not fully understood but may result in part from
the close proximity in the posterior horn of the central terminals of GVA pain fibers and
GSA spinal nerve fibers from the body surface. This is supported by the fact that pain
from a visceral origin is referred to a dermatome with which it shares the same posterior
root. This is a useful observation, often making it possible to locate the source of a
visceral pain from an observation of the surface area to which it is referred. The pain
down the inside of the left arm associated with true cardiac pain is a good example.
It is likely that separate second-order neurons relay pain information from GSA and GVA
input. If the painful stimulus to the viscera is moderate, the level of activity in the GVA
fibers is likely sufficient to stimulate only those second-order neurons which normally
relay signals from the viscera. However, if the painful stimulus increases in strength, the
increased central synaptic activity of the GVA neurons may "spill over" and raise the
central excitatory state of those second-order neurons which normally relay information
from GSA fibers of the dermatome. If the painful visceral stimulation is very strong, this
"spill over" may be sufficient to exceed the threshold of excitation for these neurons,
causing them to fire even though no painful stimulus is delivered to the general somatic
nociceptors of the dermatome. Thus the brain incorrectly projects the source of the pain
to the dermatomal area (Fig-13).

Sensory Pathways
Blood Pressure, Blood
Chemistry, and Alveolar
Stretch Detection
The walls of the aorta and
the carotid sinuses contain
special baroreceptors
(pressure receptors) which
respond to changes in
blood pressure. These
mechanoreceptors are the
peripheral endings of GVA
fibers of the
glossopharyngeal (IX) and
vagus (X) nerves. The
GVA fibers from the
carotid sinus
baroreceptors enter the
solitary tract of the
brainstem and terminate
in the vasomotor center of
the medulla (Fig-14). This
is the CNS control center
for cardiovascular activity.
The cell bodies of these
unipolar neurons are
located in the petrosal
ganglion. GVA fibers of the
vagus nerve conduct
signals from the
baroreceptors in the walls
of the aorta to the solitary
tract and on to the
vasomotor center. The cell
bodies of these unipolar
neurons are located in the
nodose ganglion.
Stretch receptors in the
alveoli of the lungs
conduct information
concerning rhythmic
alveolar inflation and
deflation over GVA X fibers
to the solitary tract and
then to the respiratory
center of the brainstem.
This route is an important
link in the Hering-Breuer
reflex, which helps to
regulate respiration.
Carotid body
chemoreceptors, sensitive
to changes in blood
PO2 and, to a lesser extent,

Fig-14

Sensory Pathways
PCO2 and pH, conduct
signals to both the
vasomotor and respiratory
centers over GVA IX nerve
fibers. GVA X fibers
conduct similar
information from the
aortic chemoreceptors to
both centers.
Chemoreceptors were
discussed elsewhere.

SPECIAL VISCERAL AFFERENT (SVA) PATHWAYS

Taste
The receptors for taste are the taste cells which produce impulses in afferent fibers in
response to chemical stimulation. They were described elsewhere. The pathways for
taste sensation are illustrated in Fig-15. Special visceral afferent (SVA) fibers of cranial
nerves VII, IX, and X conduct signals into the solitary tract of the brainstem, ultimately
terminating in the nucleus of the solitary tract on the ipsilateral side. Second-order
neurons cross over and ascend through the brainstem in the medial lemniscus to the
VPM of the thalamus. Thalamic projections to area 43 (the primary taste area) of the
postcentral gyrus complete the relay. SVA VII fibers conduct from the chemoreceptors of
taste buds on the anterior twothirds of the tongue, while SVA IX fibers conduct taste
information from buds on the posterior one-third of the tongue. SVA X fibers conduct
taste signals from those taste cells located throughout the fauces.

Fig-16
Fig-15

Sensory Pathways

Smell
The sense of smell was examined elsewhere and, once again, we will look only at the
central pathways here. The smell-sensitive cells (olfactory cells) of the olfactory
epithelium project their central processes through the cribiform plate of the ethmoid
bone, where they synapse with mitral cells. The central processes of the mitral cells pass
from the olfactory bulb through the olfactory tract, which divides into a medial and
lateral portion (Fig-16). The lateral olfactory tract terminates in the prepyriform cortex
and parts of the amygdala of the temporal lobe. These areas represent the primary
olfactory cortex. Fibers then project from here to area 28, the secondary olfactory area,
for sensory evaluation. The medial olfactory tract projects to the anterior perforated substance, the septum pellucidum, the subcallosal area, and even the contralateral olfactory
tract. Both the medial and lateral olfactory tracts contribute to the visceral reflex
pathways, causing the viscerosomatic and viscerovisceral responses described earlier.

DAMAGE TO THE SPINAL NERVES AND SPINAL CORD

After studying the motor pathways and the sensory pathways, the injuries described in
Table-1 would be expected to produce the symptoms listed.

Table-1 Symptoms of Damage to Spinal Nerves and Spinal Cord

Damage
Peripheral nerve
Posterior root

Anterior Horn

Possible cause
Symptoms associated with innervated area
of damage
Mechanical injury Loss of muscle tone. Loss of reflexes. Flaccid
paralysis. Denervation atrophy. Loss of sensation
Tabes dorsalis
Paresthesia. Intermittent sharp pains. Decreased
sensitivity to pain. Loss of reflexes. Loss of
sensation. Positive Romberg sign. High stepping
and slapping of feet.
Poliomyelitis
Loss of muscle tone. Loss of reflexes. Flaccid
paralysis. Denervation atrophy

Sensory Pathways
Lamina X (gray
matter)

Syringomyelia

Anterior horn and

lateral
corticospinal tract
Posterior and
lateral funiculi

Amyotrophic
lateral sclerosis
Subacute
combined
degeneration

Bilateral loss of pain and temperature sense only at

afflicted cord level. Sensory dissociation. No
sensory impairment below afflicted level
Muscle weakness. Muscle atrophy. Fasciculations of
hand and arm muscles. Spastic paralysis

Loss of position sense. Loss of vibratory sense.

Positive Romberg sign. Muscle weakness.
Spasticity. Hyperactive tendon reflexes. Positive
Babinski sign.
Hemisection of the Mechanical injury Brown-Sequard syndrome
spinal cord
Below cord level on injured side
Flaccid paralysis. Hyperactive tendon reflexes. Loss
of position sense. Loss of vibratory sense. Tactile
impairment
Below cord level on opposite side beginning one or
two segments below injury
Loss of pain and temperature