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164 Chapter 20<br />
2. Enkephalins are the most widely distributed and abundant opiate peptides. They are found in the<br />
highest concentration in the globus pallidus. They play a role in pain suppression.<br />
3. Dynorphins are opioid peptides that are synthesized widely throughout the CNS. They moderate<br />
pain response.<br />
E. Nonopioid Neuropeptides<br />
1. Substance P is a neuropeptide that functions in pain modulation and in signaling the intensity of<br />
noxious stimuli, more recently its role in psychologic stress has been described. It plays a role in<br />
movement disorders—substance P levels are reduced in patients with Huntington disease.<br />
2. Somatostatin (somatotropin-release–inhibiting factor). Somatostatinergic neurons from<br />
the anterior hypothalamus project their axons to the median eminence, where somatostatin enters<br />
the hypophyseal portal system and regulates the release of growth hormone and thyroidstimulating<br />
hormone. The concentration of somatostatin in the neocortex and hippocampus<br />
is significantly reduced in patients with Alzheimer disease. Striatal somatostatin levels are<br />
increased in patients with Huntington disease.<br />
F. Amino Acid Transmitters<br />
1. Inhibitory amino acid transmitters<br />
a. GABA (Figure 20-6) is the major inhibitory neurotransmitter of the brain. Purkinje, stellate,<br />
basket, and Golgi cells of the cerebellar cortex are GABA-ergic.<br />
i. GABA-ergic striatal neurons project to the globus pallidus and substantia nigra.<br />
ii. GABA-ergic pallidal neurons project to the thalamus.<br />
iii. GABA-ergic nigral neurons project to the thalamus.<br />
iv. GABA receptors (GABA-A and GABA-B) are intimately associated with benzodiazepinebinding<br />
sites. Benzodiazepines enhance GABA activity.<br />
b. Glycine is a major inhibitory neurotransmitter of the spinal cord, brainstem, and retina. It is<br />
used by the Renshaw cells of the spinal cord.<br />
2. Excitatory amino acid transmitters<br />
a. Glutamate (Figure 20-7) is the major excitatory neurotransmitter of the CNS. Neocortical<br />
glutamatergic neurons project to the striatum, subthalamic nucleus, and thalamus.<br />
i. Glutamate is the transmitter of the cerebellar granule cells, of nonnociceptive, large, primary<br />
afferent fibers that enter the spinal cord and brainstem, and of the corticonuclear and corticospinal<br />
tracts.<br />
Figure 20-6 Distribution of γ-aminobutyric acid (GABA)-containing neurons and their projections. GABA-ergic neurons<br />
are the major inhibitory cells of the central nervous system. GABA local circuit neurons are found in the neocortex,<br />
hippocampal formation, and cerebellar cortex (Purkinje cells). Striatal GABA-ergic neurons project to the thalamus and<br />
subthalamic nucleus (not shown).