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Ganong's Review of Medical Physiology, 23rd Edition

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Amygdaloid<br />

nucleus<br />

Caudate nucleus<br />

Globus<br />

pallidus<br />

Caudate nucleus<br />

Lateral view<br />

Putamen<br />

Thalamus<br />

Putamen and<br />

globus pallidus<br />

Horizontal section<br />

Thalamus<br />

Tail <strong>of</strong><br />

caudate nucleus<br />

The main feature <strong>of</strong> the connections <strong>of</strong> the basal ganglia is<br />

that the cerebral cortex projects to the striatum, the striatum<br />

to GPi, GPi to the thalamus, and the thalamus back to the cortex,<br />

completing a loop. The output from GPi to the thalamus<br />

is inhibitory, whereas the output from the thalamus to the<br />

cerebral cortex is excitatory.<br />

CHAPTER 16 Control <strong>of</strong> Posture & Movement 251<br />

Substantia<br />

nigra<br />

Amygdala<br />

FIGURE 16–9 The basal ganglia. The basal ganglia are composed <strong>of</strong> the caudate nucleus, putamen, and globus pallidus and the functionally<br />

related subthalamic nucleus and substantia nigra. The frontal (coronal) section shows the location <strong>of</strong> the basal ganglia in relation to surrounding<br />

structures.<br />

Globus<br />

pallidus, ES<br />

GABA<br />

Glu<br />

Subthalamic<br />

nucleus<br />

Brain stem<br />

and<br />

spinal cord<br />

Glu<br />

GABA<br />

Internal capsule<br />

GABA<br />

Cerebral<br />

cortex<br />

Glu<br />

Striatum<br />

(acetylcholine)<br />

Globus<br />

pallidus, IS<br />

GABA<br />

GABA GABA GABA<br />

PPN<br />

Thalamus<br />

Thalamus<br />

Internal<br />

capsule<br />

FIGURE 16–10 Diagrammatic representation <strong>of</strong> the principal<br />

connections <strong>of</strong> the basal ganglia. Solid lines indicate excitatory pathways,<br />

dashed lines inhibitory pathways. The transmitters are indicated<br />

in the pathways, where they are known. Glu, glutamate; DA, dopamine.<br />

Acetylcholine is the transmitter produced by interneurons in the striatum.<br />

SNPR, substantia nigra, pars reticulata; SNPC, substantia nigra, pars<br />

compacta; ES, external segment; IS, internal segment; PPN, pedunculopontine<br />

nuclei. The subthalamic nucleus also projects to the pars compacta<br />

<strong>of</strong> the substantia nigra; this pathway has been omitted for clarity.<br />

DA<br />

SNPR SNPC<br />

Frontal section<br />

The striatum is made up <strong>of</strong> two parts that differ histologically:<br />

a unique mosaic <strong>of</strong> patches or striosomes (an area with<br />

little acetylcholinesterase) and a matrix (an area high in acetylcholinesterase).<br />

The neurons <strong>of</strong> the corticostriate projection<br />

that originate in the deep portion <strong>of</strong> layer V <strong>of</strong> the cortex<br />

terminate in the patches, whereas the neurons that originate<br />

in layers II and III and the superficial part <strong>of</strong> layer V end primarily<br />

in the matrix. Neurons with their cell bodies in patches<br />

project in large part to dopaminergic neurons in the substantia<br />

nigra pars compacta, whereas many <strong>of</strong> the neurons with<br />

their cell bodies in the matrix project to GABAergic neurons<br />

in the substantia nigra pars reticulata.<br />

FUNCTION<br />

Lateral ventricle<br />

Caudate nucleus<br />

Putamen<br />

Globus pallidus:<br />

External segment<br />

Internal segment<br />

Subthalamic<br />

nucleus<br />

Neurons in the basal ganglia, like those in the lateral portions<br />

<strong>of</strong> the cerebellar hemispheres, discharge before movements<br />

begin. This observation, plus careful analysis <strong>of</strong> the effects <strong>of</strong><br />

diseases <strong>of</strong> the basal ganglion in humans and the effects <strong>of</strong><br />

drugs that destroy dopaminergic neurons in animals, have led<br />

to the idea that the basal ganglia are involved in the planning<br />

and programming <strong>of</strong> movement or, more broadly, in the processes<br />

by which an abstract thought is converted into voluntary<br />

action (Figure 16–1). They influence the motor cortex via<br />

the thalamus, and the corticospinal pathways provide the final<br />

common pathway to motor neurons. In addition, GPi projects<br />

to nuclei in the brain stem, and from there to motor neurons<br />

in the brain stem and spinal cord. The basal ganglia, particularly

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