


The two types of pain fibers
enter different layers of the dorsal horn.
Ad
fibers enter the
posterior marginalis and the nucleus proprius, and synapse
on a second set of neurons. These are the secondary
afferents (purple, below) which will carry the signal to
the thalamus. The secondary afferents from both layers
cross to the opposite side of the spinal cord and ascend in
a tract called (logically) the
spinothalamic tract. Tracts are always labeled
from beginning to end.

The C fibers enter the
substantia gelatinosa and synapse, but they do not
synapse on secondary afferents. Instead they synapse
on
interneurons - neurons which do not project
out of the immediate area. The interneurons must carry the
signal to the secondary afferents in either the posterior
marginalis or the nucleus proprius.

The spinothalamic tract
ascends the entire length of the cord as shown above,
and the entire brainstem, staying in about the same
location all the way up. Below are representative
slides showing the tract in the medulla and midbrain.
Notice that by midbrain the spinothalamic tract
appears to be continuous with the medial lemniscus.
They will enter the VPL of the thalamus
together.

The spinothalamic system
enters the VPL, synapses, and is finally carried to
cortex by the thalamocortical neurons. Here is a
schematic of the entire pathway:

E.
Pain control:
It has been recognized for
centuries that
opium and
related compounds (such as
morphine)
are powerful analgesics. Several decades ago scientists
hunted down the opiate receptor which was responsible for
the potent effects. They then reasoned that if there was
such a receptor in the body, maybe the body used its own
endogenous form of opium to control pain. (It has also been
recognized for centuries that under certain circumstances,
i.e. the heat of battle, a serious wound may not cause
pain.) This hypothetical compound was named
"endorphin",
from endogenous-morphine. Soon after, an entire class of
peptide neurotransmitters was discovered that interacted
with the opiate receptor, and now includes endorphins,
enkephalins, and dynorphins. Synthetic, exogenous forms of
these compounds continue to be discovered, prescribed, and
abused, and are classed under the general term,
"narcotics".
There are opiate receptors
throughout the central nervous system. In the dorsal horn,
they are located on the terminals of the primary afferents,
as well as on the cell bodies of the secondary afferents.
Opiate interneurons in the spinal cord can be activated by
descending projections from the brainstem (especially the
raphe nuclei and periaqueductal grey), and can block pain
transmission at two sites. 1) They can prevent the primary
afferent from passing on its signal by blocking
neurotransmitter release, and 2) they can inhibit the
secondary afferent so it does not send the signal up the
spinothalamic tract.

F.
The proprioceptive system:
The proprioceptive system
arises from primarily the Aa afferents entering the
spinal cord. These are the afferents from muscle
spindles, Golgi tendon organs, and joint receptors.
The axons travel for a little while with the
discriminative touch system, in the posterior columns.
Within a few segments, however, the proprioceptive
information slips out of the dorsal white matter and
synapses. After synapsing it ascends without
crossing to the cerebellum.
Exactly
where the axons synapse depends upon whether they
originated in the legs or the arms. Leg fibers enter
the cord at sacral or lumbar levels, ascend to the
upper lumbar segments, and synapse in a medial nucleus
called
Clarke's nucleus (or nucleus dorsalis). The
secondary afferents then enter the
dorsal
spinocerebellar tract on the lateral edge of the
cord.

Fibers from the arm enter at
cervical levels and ascend to the caudal medulla. Once
there they synapse in a nucleus called the
external cuneate (or lateral cuneate) nucleus,
and the secondary axons join the leg information in the
dorsal spinocerebellar tract.

The spinocerebellar tract
stays on the lateral margin of the brainstem all the
way up the medulla. Just before reaching the pons, it
is joined by a large projection from the
inferior olive. These axons together make up
the bulk of the
inferior cerebellar peduncle, which grows right out of the
lateral medulla and enters the
cerebellum.

The figures above outline
the course of the dorsal spinocerebellar tract.
Surely, there must be a ventral spinocerebellar tract?
Naturally, there is, and it travels in approximately
the same place - the lateral margin of the spinal
cord, just ventral to the dorsal spinocerebellar
tract. The two cannot be distinguished in a normal
myelin stain. The ventral spinocerebellar tract seems
to defy the ipsilaterality of the cerebellum, because
the fibers entering it in the spinal cord actually
cross on their way into the tract. However, they
(somewhat inefficiently) cross back before entering
the cerebellum. Therefore the cerebellum still gets
information from the ipsilateral
body.
A note
about generalizations:
There is actually a fair
amount of mixing that goes on between the tracts. Some
light touch information travels in the spinothalamic
tract, so that lesioning the dorsal columns will not
completely knock out touch and pressure sensation.
Some proprioception also travels in the dorsal
columns, and follows the medial lemniscus all the way
to the cortex, so there is conscious awareness of body
position and movement. The pain and temperature
system, although it does ascend to somatosensory
cortex, also has multiple targets in the brainstem and
other areas.
For a more interactive
tutorial on the anatomy, and for stunning three
dimensional pictures of these pathways, try the
Digital Anatomist in the "Other links"
section.