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skeletal muscles. The basal ganglia refine action signals from the cortex, thereby ensuring
that an appropriate motor plan is communicated to the muscles. Unlike the pyramidal
pathway, the basal ganglia process information indirectly in a set of loops, whereby they
receive input from the cortex and return it to the cortex via the thalamus. In that way the basal
ganglia modify the timing and amount of activity that leaves the cortex and travels down the
pyramidal pathway, amplifying activity that leads to a positive outcome and suppressing
activity that leads to a deleterious outcome in a particular situation.
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Much knowledge about the role of the basal ganglia in brain function has come from the
study of disorders that affect the different nuclei. Typically, such disorders lead to difficulty
with initiating wanted movements (as generally seen in Parkinson disease) or with
suppressing unwanted movements (as seen in Huntington disease).
The major input nucleus of the basal ganglia is the striatum (collectively including the dorsal
and ventral divisions), which receives information from almost all areas of the cortex. The
dorsal striatum (upper region of the striatum) receives information from areas below the
cortex (e.g., the midbrain) via the thalamus. In the motor circuit the subthalamic nucleus
serves as an input nucleus, receiving information from the cortex and thalamus and
influencing the conventional route of basal ganglia outflow from the striatum to the output
nuclei of the thalamus. The output nuclei of the basal ganglia are the globus pallidus internus
and substantia nigra pars reticulata in the motor pathway and the ventral pallidum in the
limbic pathway. Information that exits the basal ganglia goes to the thalamus, primarily the
ventroanterior and ventromedial motor thalamic nuclei for the motor pathway and the
mediodorsal thalamic nucleus for the limbic pathway, and then is sent back to the appropriate
part of the cortex.
Neurochemicals
The majority of basal ganglia nuclei have projection neurons (neurons with axons that extend
into adjacentbrain areas) that utilize the inhibitory neurotransmitter gamma-aminobutyric
acid (GABA). As a result, inhibitory signals form the basis of most communication between
nuclei in the basal ganglia. Exceptions include the excitatory glutamate-releasing projections
of the subthalamic nucleus and the dopamine-releasing projection neurons from the
substantia nigra pars compacta.
The striatum, which serves as a gateway for the regulation of signals through the basal
ganglia during the learning of actions and the selection of desirable actions, has the most-
complex signaling architecture. In addition to receiving vast external excitatory input from
the cortex and thalamus, it also contains several types of interneurons (neurons that connect
sensory and motor circuits) and some of the highest levels in the brain of the neurochemicals
dopamine and acetylcholine. Collectively, these substances modulate the way in which
excitatory inputs are processed and contribute to the final output from the striatum.
Cell Physiology
The brain encodes and transmits information between areas in the form of electrical impulses
called action potentials. The processing and relaying of information in the basal ganglia are
complex, because the majority of neurons release GABA when they fire action potentials,
generally inhibiting the activity of cells in the target areas. Therefore, a basic operating
principle of information progression through the basal ganglia is the removal of the net
inhibition imposed by output nuclei onto target areas in the thalamus and cortex, a process
known as disinhibition. The final behavioral outcome depends on the timing and
spatial dynamics of firing events in single neurons and groups of neurons (local networks) as
well as across parallel pathways (large networks).
The importance of the basal ganglia in generating movements is evident from the rate and
pattern of action potentials fired in neurons during the preparation for and execution of
movements. The majority of neurons alter their activity after the movement has started,
which supports the idea that the basal ganglia are able to fine-tune movements. Some neurons
in the basal ganglia, however, have precise roles in learning and the cueing of movement. For
instance, neurons in the striatum that manufacture acetylcholine show a dramatic pause in
their firing when a sensory signal (e.g., a flash of light or unusual sound) is associated with a
meaningful action (e.g., sitting or running). Such signals conversely cause dopamine neurons
in the substantia nigra pars compacta and ventral tegmental area to fire faster for a few
hundredths of a second, thereby releasing pulses of dopamine into the striatum. Together, the
timing of acetylcholine and dopamine release teaches the striatum which signals to pay
attention to (e.g., signals that lead to a rewarding outcome) and allows it to learn which action
recently performed led to the appearance of these signals. This results in the reinforcement of
specific pathways through the striatum, ensuring that desirable actions reoccur more
frequently in the future. Through this process, for example, a dog learns that a whistle from
its owner will lead to a treat after it performs the requested action of sitting.
Reinforcement occurs at the cellular level by strengthening synaptic inputs from the cortex
onto cells in the striatum through a mechanism called synaptic plasticity. Dopamine plays a
key role in this process and is essential for both strengthening synaptic inputs as well as
weakening synaptic inputs that code for unwanted and undesirable motor plans. Thus,
dopamine neurons act as gatekeepers, controlling which messages progress from the striatum
to other basal ganglia nuclei during the action-selection process. Furthermore, through the
activity of dopamine neurons, the basal ganglia also provide the motivation to perform
behaviours that are required to explore, interact with, and learn from one’s environment.
human nervous system: Basal ganglia
Deep within the cerebral hemispheres, large gray masses of nerve cells, called nuclei, form
components…
human nervous system: Basal ganglia
Most of what is known about the contribution of the basal ganglia has been obtained from
studying abnormal…
Parkinson disease is a progressive disorder caused by degeneration of the cells of the substantia
nigra…
…lobe), the archipallium, and the basal nuclei. All three areas receive olfactory stimuli and
discharge…
Huntington disease
…death of neurons in the basal ganglia, a pair of nerve clusters deep within the brain that control
movement.…
Assorted References
major reference
In human nervous system: Basal ganglia
In human nervous system: Basal ganglia
Huntington disease
In Huntington disease
neurological disease
In nervous system disease: Basal ganglia and thalamus
In nervous system disease: The basal ganglia
vertebrate nervous systems
In nervous system: Dominance of the cerebrum
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