154 Section III Organization of the Central Nervous System

CHAPTER SUMMARY TABLE

Spinal Cord Tracts
Tract Receptors Course and Distributionb Functions Effects of Lesions
Long Ascending Spinal Tracts

Fasciculus Meissners Exists at all levels of spinal cord; located medial to Carries information about Loss of sensations of fine
gracilisa (touch), fasciculus cuneatus, central processes of first- fine touch, pressure, touch, pressure, vibra-
Merkels (pres- order neurons ascend ipsilaterally and terminate vibration sensation, and tion sensation, and
sure), Pacinian on second-order neurons located in ipsilateral conscious propriocep- conscious propriocep-
(vibration), and medullary nucleus gracilis; axons of second-order tion (sense of position tion from lower body
joint receptors neurons cross in midline (internal arcuate fibers) and movement) from and limbs ipsilateral
(kinesthesia to form medial lemniscus and ascend to synapse the lower body and to lesion; ataxia due
position and on third-order neurons located in contralateral limbs (sacral, lumbar to loss of conscious
movement) thalamic ventral posterolateral nucleus; axons of and T6T12 region) to proprioception from
third-order neurons terminate in medial aspect of brainstem lower limbs
sensorimotor cortex

Fasciculus Meissners Exists at T6 thoracic levels and cervical levels; located Carries information about Loss of sensations of
cuneatusa (touch), lateral to fasciculus gracilis; central processes fine touch, pressure, fine touch, pressure,
Merkels (pres- of first-order neurons ascend ipsilaterally and vibration sensation, and vibration sensation, and
sure), Pacinian terminate on second-order neurons located in conscious propriocep- conscious proprioception
(vibration), and ipsilateral medullary nucleus cuneatus; axons of tion (sense of position from upper body and
joint receptors second-order neurons project to third-order neu- and movement) from limbs ipsilateral to lesion
(kinesthesia rons located in contralateral thalamic ventral pos- upper body and limbs
position and terolateral nucleus; axons of third-order neurons to brainstem
movement) terminate in lateral aspect of sensorimotor cortex

Dorsal Muscle spindles, Central processes of first-order neurons project Transmits signals from Likely gait ataxia
(posterior) Golgi tendon to neurons in nucleus dorsalis of Clarke (C8 to muscle spindle and Golgi
spino- organs in lower L2); axons of these neurons ascend ipsilaterally, tendon afferents in lower
cerebellar limbs reach inferior cerebellar peduncle in medulla, limb to cerebellum, which
tract and terminate ipsilaterally in cerebellar vermis of coordinates noncon-
anterior lobe scious proprioception

Ventral Golgi tendon Central processes of first-order neurons project to Transmits information Unknown
(anterior) organs in lower second-order neurons in dorsal horn; axons of most from Golgi tendon
spino- limbs second-order neurons cross to contralateral lateral organs located in lower
cerebellar funiculus and ascend to pons where they join supe- limb to cerebellum
tract rior cerebellar peduncle, cross again to other side,
and terminate in cerebellar vermis of anterior lobe

Cuneo- Muscle spindles, Central processes of first-order neurons ascend ipsilater- Transmits information Loss of nonconscious
cerebellar Golgi tendon ally in fasciculus cuneatus and synapse on neurons in about muscle spindle proprioception and
tract organs in upper accessory cuneate nucleus; neurons in this nucleus and Golgi tendon coordination of upper
limbs give rise to cuneocerebellar tract, which terminates afferents in upper limb limbs ipsilateral to
ipsilaterally in cerebellar vermis of anterior lobe to cerebellum lesion

Rostral spino- Golgi tendon Same as in ventral spinocerebellar tract except that Transmits information Unknown
cerebellar organs in upper afferents are from upper limb, tract is uncrossed, and from Golgi tendon
tract limbs it enters cerebellum via inferior cerebellar peduncle organs located in upper
(not known if present in humans) limb to cerebellum

Spinothalamic Nociceptors: Free Direct: Mediates discriminative Cervical Lesion: Complete

tractc nerve endings Neospinothalamic Tract: Neurons that give rise component of pain loss of pain, temperature,
mediating to this tract arise mainly from proper sensory sensation and simple tactile sensa-
mechanical, nucleus (lamina III and IV); axons of these tions on contralateral side
thermal, and neurons cross obliquely to enter contralateral of body (upper and lower
polymodal white matter, ascend in lateral funiculus, and limbs and trunk)
sensations synapse on third-order neurons located primar- Lumbar Lesion: Loss of
ily in ventral posterolateral nucleus of thalamus pain, temperature, and
that projects to primary somatosensory cortex in simple tactile sensa-
postcentral gyrus tions in contralateral
lower limb and trunk
Tract Receptors Course and Distributionb Functions Effects of Lesions
Long Ascending Spinal Tracts

Indirect: Mediate arousal-emotional Lesions of these tracts

Paleospinothalamic Tract: Neurons of this pathway components of pain are believed to cause
are located in dorsal horn and intermediate gray sensation a loss of arousal-
matter; axons of these neurons ascend bilaterally, emotional components
make several synapses in reticular formation of of pain
medulla, pons, and midbrain, and finally project
to midline and intralaminar thalamic nuclei, which
then project in a diffuse manner to cerebral cortex.
Spinoreticular Tract: Neurons of this pathway are also
located in dorsal horn and intermediate gray mat-
ter; their axons ascend bilaterally and terminate
on neurons located in medullary and pontine
reticular formation; these neurons activate cere-
bral cortex through secondary and tertiary projec-
tions via midline and intralaminar thalamic nuclei.
Spinomesencephalic Tract: Neurons of this pathway are
also located in dorsal horn and intermediate gray
matter; axons of these neurons ascend to midbrain
where they terminate in PAG; activation of neurons
in PAG results in inhibition of pain sensation at spi-
nal level via indirect descending projections; sensory
information carried by this tract is also transmitted
to amygdala via parabrachial nuclei
Long Descending Spinal Tracts

Corticospinal Cerebral cortex Arises from cerebral cortex, passes through medul- Controls voluntary Voluntary control of con-
tract lary pyramids, and terminates in spinal cord; in movements of both tralateral upper and
cortex, cells functionally associated with arm and contralateral upper and lower limbs is lost when
leg are located in lateral convexity and medial wall lower limbs corticospinal tract is
of hemisphere, respectively (cortical homunculus); damaged; symptoms of
axons arising from cortex converge in corona damage to corticospinal
radiata and descend through internal capsule, tract (i.e., loss of volun-
crus cerebri in midbrain, pons, and medulla; a tary movement, spas-
majority (about 90%) of fibers cross to contralat- ticity, increased deep
eral side at juncture of medulla and spinal cord tendon reflexes, loss of
(pyramidal decussation), forming lateral cortico- superficial reflexes, and
spinal tract, which descends to all levels of spinal Babinski sign) comprise
cord and terminates in spinal gray matter of both an upper motor neuron
dorsal and ventral horns; remaining uncrossed fi- paralysis; symptoms
bers (anterior corticospinal tract) descend through of lower motor neuron
spinal cord and cross over at different segmental paralysis include loss
levels to synapse with anterior horn cells on con- of muscle tone, atrophy
tralateral side; pyramidal decussation forms ana- of muscles, and loss of
tomical basis for voluntary motor control of one all reflex and voluntary
half of body by contralateral cerebral hemisphere movement

Rubrospinal Red nucleus Axons of red nucleus neurons cross midline in ventral Function of this tract is to Although effects of lesions
tract (located in the midbrain (ventral tegmental decussation) and de- facilitate flexor motor restricted to red
rostral half of scend to contralateral spinal cord; fibers in rubro- neurons and inhibit nucleus are not known
the midbrain spinal tract are somatotopically arranged; cervical extensor motor neurons in humans, lesions of
tegmentum) spinal segments receive fibers from dorsal part midbrain tegmentum
of red nucleus, which receives inputs from upper including red nucleus
limb region of sensorimotor cortex; lumbosacral are reported to elicit
spinal segments receive fibers from ventral half of contralateral motor dis-
red nucleus, which receives inputs from lower limb turbances (tremor, ataxia,
region of sensorimotor cortex; fibers of rubrospinal and choreiform activity),
tract terminate on interneurons that, in turn, proj- possibly because of
ect to dorsal aspect of ventral (motor) horn cells; in involvement of axons
humans most of efferent fibers emerging from red arising from basal gan-
nucleus terminate in inferior olive and cervical cord glia or cerebellum
(continued on page 156)
156 Section III Organization of the Central Nervous System

CHAPTER SUMMARY TABLE

Spinal Cord Tracts (continued)
Tract Receptors Course and Distributionb Functions Effects of Lesions
Long Descending Spinal Tracts

Tectospinal Superior colliculus Axons of neurons in superior colliculus terminate in May direct head move- Not established
tract upper cervical segments ments in response to
visual and auditory
stimuli

Lateral ves- Lateral vestibular Fibers in this tract are uncrossed, descend entire Facilitates ipsilateral Disturbances in main-
tibulospinal nucleus (border length of spinal cord, and terminate on inter- extensor alpha-motor taining posture and
tract of the pons and neurons that activate motor neurons innervating neurons and associated balance; effects of
medulla) extensor muscles of trunk and ipsilateral limb gamma motor neurons; lesions limited to
main function is to lateral vestibulospinal
control muscles that tract have not been
maintain upright posture described in humans
and balance

Medial ves- Ipsilateral and Descends in ventral funiculus of cervical spinal cord Main function of this tract Possible disturbances in
tibulospinal contralateral and terminates in ipsilateral ventral horn is to adjust position positioning of head
tract medial vestibu- of head in response when position of body
lar nuclei to changes in posture is affected
(such as keeping head
stable while walking)

Reticulospinal Medulla: Nucleus Projects bilaterally to all levels of spinal Inhibits extensor spinal Believed to contribute
tracts reticularis cord; this tract is called medullary (lateral) reflex to spasticity in upper
gigantocellularis reticulospinal tract motor neuron paralysis

Pons: Nucleus Projects ipsilaterally to entire spinal cord; this tract Facilitates extensor spinal Not established; possible
reticularis pontis is called pontine (medial) reticulospinal tract reflexes loss of muscle tone of
caudalis and extensor muscles
oralis

Ventrolateral Projects to IML of thoracolumbar cord Excites sympathetic
medulla preganglionic neurons
in IML, which provide
sympathetic innervation
to visceral organs
Not established; presumed
to contribute to Horners
syndrome

PAG Enkephalinergic neurons located in midbrain Modulates activity of pain Not established; possible
PAG project to serotonergic neurons located in impulses that ascend in disturbance of pain
nucleus raphe magnus of medulla (first limb); spinothalamic system sensation
second limb of pathway consists of projections
from serotonergic raphe magnus neurons to
enkephalinergic interneurons in dorsal horn of
spinal cord, which, in turn, synapse upon primary
afferent pain fibers
 Chapter 8 The Spinal Cord 157

Tract Receptors Course and Distributionb Functions Effects of Lesions

Long Descending Spinal Tracts

MLF Medial vestibular Descending fibers in MLF project primarily to Descending MLF fibers Although damage to
nucleus, reticu- ipsilateral upper cervical spinal cord segments monosynaptically descending fibers of
lar formation, inhibit motor neurons MLF is presumed to
and superior located in upper cervi- disrupt control of posi-
colliculus cal spinal segments tion of head in response
(tectospinal and control position of to inputs from labyrinth
fibers) head in response to and vestibular appara-
excitation by labyrinth tus, these symptoms
of vestibular apparatus have not been clinically
established; damage
to ascending fibers
in MLF at the level of
brainstem produces
internuclear ophthalmo-
plegias; lesions of MLF
are common in patients
with multiple sclerosis
in which symptoms
include nystagmus
and occasionally dip-
lopia in ipsilateral eye;
these symptoms may
be due to damage to
projections from lateral,
superior, and medial
vestibular nuclei to
oculomotor, abducens,
and trochlear cranial
nerve nuclei

IML, intermediolateral cell column; MLF, medial longitudinal fasciculus; PAG, periaqueductal midbrain gray.
a
Fasciculi gracilis and cuneatus together form the dorsal column of the spinal cord.
b
First-order neuron of all ascending tracts are located in the dorsal root ganglia.
c
Direct (neospinothalamic) and indirect (paleospinothalamic, spinoreticular, and spinomesencephalic) tracts comprising
the spinothalamic tract are collectively known as the anterolateral system of ascending tracts.