Edmund
Rolls - Brain and Emotion |
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Prefrontal cortex has undergone great development in primates, and the orbitofrontal cortex is very little developed in rodents, yet is it is one of the major
brain areas involved in emotion and motivation and primates including humans. |
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A reward is something for which an animal will work. |
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A punishment is something an animal of will work to escape or avoid. |
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In classical
conditioning, the taste of food (the unconditioned stimulus) might
elicit an unconditioned response of salivation, and if the sight of the food
is paierd with the taste, then the sight of that food would by learning come
to produce salivation, and would become a conditioned
stimulus for the conditioned
response of salivation. |
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Pairing the
sight of an object
with a reward or punishment can lead to stimulus-reinforcement association
learning. |
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Having reward and punishment systems is a solution that evolution has developed to produce
appropriate behavior. |
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Motivational and emotional behavior are the types of behavior in which rewards and punishers operate. |
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A distinction is that motivated behavior often refers to behavior where the initiating
stimulus is an internal, whereas emotional behavior often refers to behavior where the initiating
stimulus is an external. |
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The brain works in motivation and emotion to result in natural selection operating to select genes which optimize our behavior by building into us the appropriate reward and punishment systems. |
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Responses of hypothalamic
neurons in the primate become associated with the sight of food as a result of learning. |
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There are neurons in the
hypothalamus that can respond to the taste (and/or sight) of foods but not of
non-foods. |
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An important food taste that
appears to be different from the taste produced by sweet, salt, bitter, or
sour is the taste of protein. |
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The umami taste is common to a diversity of
food sources including fish, meats, mushrooms,
cheese, and some vegetables such as tomatoes. |
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Monosodium glutamate (MSG) is an example of umami stimuli. |
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The primate amygdala contains a population of neurons specialized to respond to faces, and damage to the
amygdala can alter the ability to discriminate between different facial
expressions. |
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Connections to the Amygdala |
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The amygdala is a subcortical
region in the anterior part of the temporal lobe. |
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The amygdala receives massive projections from the overlying temporal lobe
cortex. |
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Inputs to the amygdala come from the inferior temporal visual cortex, the
superior temporal auditory cortex, the
cortex of the temporal pole, and the
cortex of the superior temporal sulcus. |
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Inputs to the amygdala come from the higher stages of sensory processing in the visual and auditory modalities, and not from early cortical processing
areas. |
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The amygdala also receives inputs
that are potentially about primary reinforces, e.g. taste inputs (from the secondary taste
cortex, via connections from the orbitofrontal cortex to the
amygdala), and somatosensory inputs, potentially about the rewarding
or painful aspects of touch (from the somatosensory
cortex via the insula). |
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The amygdala also receives
projections from the posterior orbitofrontal cortex. |
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It has been suggested that the
function of backprojections
from the amygdala to the cortex include the guidance of information representation and storage in the neocortex, and the recall of the information. |
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The amygdala has output projections to the entorhinal cortex, which provides a major input to the hippocampus and dentate gyrus, and to the ventral subiculum, which provides a major output to the hippocampus. |
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Anatomical connections of the amygdala indicate that it is strategically placed to receive highly processed information
from the cortex and to influence motor systems,
autonomic systems, and some
of the cortical areas from which it receives
inputs, and other limbic areas. |
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Orbitofrontal Cortex |
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Prefrontal cortex is a region of cortex that receives projections from the medial dorsal nucleus of the thalamus and is situated in front of the motor and premotor cortices. |
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Researchers discovered a taste area in the lateral part of the orbitofrontal
cortex and showed that this was secondary taste cortex that receives a major projection
from the primary taste
cortex. |
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Convergence
of taste and olfactory inputs in the orbitofrontal cortex, forming the
representation of flavor. |
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There is a major
visual input to many
neurons in the orbitofrontal cortex, and what is represented by these
neurons in many cases is the reinforcement association of visual stimuli. |
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Visual input
to the orbitofrontal
cortex is from the ventral
temporal lobe visual stream concerned with 'what' object is being seen. |
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Representation of faces in the orbitofrontal cortex. |
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There is a population of orbitofrontal face-selective neurons
that respond in many ways similarly to those in
the temporal cortical visual areas. |
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Orbitofrontal face responsive
neurons tend to respond with
longer latencies (140-200
ms typically) than temporal
lobe neurons (80-100
ms). They also convey information about which face is being seen, by having different responses to different faces. |
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Some of the orbitofrontal
face-selective neurons are
responsive to face gesture or movement. |
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The significance of the face gesture or movement neurons in orbitofrontal cortex is likely to be that faces convey
information that is important in social reinforcement, both by conveying face expression and by
encoding the identity of an individual, both types of information important in social situations. |
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Neurophysiological evidence
suggests that one function implemented by the orbitofrontal
cortex is rapid stimulus-reinforcer association learning, and the correction of these
associations when reinforcement
contingencies in the environment
change. |
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Orbitofrontal cortex has the necessary representation
of primary reinforces,
such as taste, and also somatosensory sensory inputs. |
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Olfactory stimulus can also be conditioned via stimulus-reinforcer
association learning in the orbitofrontal cortex, although the
learning is slower
than for visual. |
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It is likely that auditory stimuli can be associated with primary reinforcers
in the orbitofrontal cortex. |
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Orbitofrontal cortex neurons which can detect non-reward in a context specific manner, are likely to be used in behavioral
extinction and reversal. |
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It is via the striatal
route that the orbitofrontal
cortex may directly
influence behavior when the orbitofrontal cortex is decoding reinforcement contingencies in the environment. |
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The amygdala the is concerned with some of the same functions as the orbitofrontal cortex and receives similar inputs. |
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There is some evidence that the amygdala may function less effectively in the very rapid learning and reversal of stimulus-reinforcement associations, as indicated by the greater difficulty in obtaining reversals from amygdala neurons. |
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In primates, the necessity of a very rapid
stimulus-reinforcement reevaluation and the
development of powerful cortical learning systems may result in the orbitofrontal
cortex effectively taking
over this aspect of amygdala
functions. |
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Active presynaptic inputs and little postsynaptic activation is the condition for homosynaptic long
term depression. |
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Wisconsin card sorting task (in which cards are to be sorted according to the color, shape, or number of items on each card,
depending on whether the examinor says 'right' or 'wrong' to each placement). |
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Anterior cingulate cortex is connected to the medial orbitofrontal areas, parts
of lateral orbitofrontal area 12, the amygdala (which projects strongly to cingulate subgenual area 25), and
the temporal pole cortex,
and also receives somatosensory inputs from the insula and other somatosensory cortical
areas. |
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Anterior cingulate cortex has output projections to the para-aqueductal gray in the midbrain (which is implicated in pain processing), to the nucleus of
the solitary tract and dorsal motor nucleus of the vagus (through which autonomic effects can be elicited), and to the
ventral striatum and caudate nucleus (through which
behavioral responses could be produced). |
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The orbitofrontal
cortex is involved in the execution of behavioral responses
when they are computed by reward or punisher association learning, a function for which the orbitofrontal cortex is
specialized. |
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Orbitofrontal cortex is specialized for reward or punisher association learning in terms of representations of primary (unlearned) reinforcers, and in rapid learning and
readjusting associations of stimuli with these primary reinforces. |
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Adapted value of the cortical strobe provided by the basal magnocellular neurons of the amygdala may be that it facilitates memory storage especially
when significant (e.g.
reinforcing) environmental stimuli are detected. |
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Memory storage is likely to be conserved (new
memories are less
likely to be laid
down) when significant
environmental stimuli are not present. |
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Actual memory stored are
determined by the act is subset of the thousands of cortical afferents own a
strongly activated cortical neuron. |
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Basal forebrain magnocellular
neurons, when activated, increased the probability that a memory will be
stored. |
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Impairment
of the normal operation of the basal forebrain
magnocellular neurons would be expected to interfere with normal memory, and
this interference could contribute to the memory disorder in Alzheimer's disease. |
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There is typically an onset of the neuronal response at 80-100 ms after the stimulus,
followed within 50 ms
by the highest firing rate. |
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When cells are responding fast (e.g. 100
spikes per second) to a visual
stimulus. |
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Source of noradrenergic projection to the neocortex is the locus coeruleus in the pons. A few thousand of these neurons innervate the whole of the cerebral cortex, as well is the amygdala and other structures. |
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To the extent that noradrenergic neurons are involved
in memory, it is
likely they would have a modulatory role on cell excitability, which would influence the extent to which voltage-dependent NMDA receptors
are activated, and
thus the likelihood that information carried on specific afferents would be stored. |
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From research studies on animals
it has been found that the lateral hypothalamus,
orbitofrontal cortex, amygdala, nucleus accumbens,
and ventral tegmental area are involved in the neurophysiology
of reward. |
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In the neurophysiology
of reward, there is a highly
interconnected set of structures, whereby
activity in any one will tend to stimulate neurons in the others. |
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Secondary taste cortex is in the orbitofrontal cortex. |
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Representation of taste in the secondary taste cortex is of the reward value of taste. |
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Reward value
of olfactory stimuli
is represented in the secondary and tertiary cortical olfactory
areas in the primate orbitofrontal
cortex. |
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Reward value
of visual stimuli such is the sight of food is represented in the primate orbitofrontal cortex, and the learning of the representation of which visual stimuli are rewarding is built in the
orbitofrontal cortex by visual-to-taste a reward learning. |
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Addictive drugs amphetamine and cocaine produce their reward by
acting on the dopaminergic projections to the nucleus accumbens. |
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Dopamine
appears to modulate transmission which links rewards systems and
structures such as the amygdala and orbitofrontal cortex via their connections to the ventral
striatum to behavioral
response selection and execution systems. |
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Two catecholamines in the brain that we are concerned with are dopamine and noradrenaline (which is also known
by its Greek name norepinephrine). |
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After dopamine is released into the synapse, and some of it activates the
postsynaptic receptors, the remaining dopamine is removed from the synapse quickly
by a number of mechanisms. |
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One mechanism for removing dopamine from the synapse is reuptake into the presynaptic terminal. |
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Another mechanism for removing dopamine from the synapse is by monoamine oxidase (MAO), which destroys the dopamine. |
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Yet another mechanism for removing dopamine from the synapse
is diffusion out of
the synaptic cleft. |
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General connectivity of the basal ganglia is for cortical or limbic inputs to reach the striatum, which then projects to the globus pallidus and substantia nigra, which in turn
project via the thalamus
back to the cerebral cortex. Within this overall scheme, there is a set of it least
partially segregated parallel
processing streams. |
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Limbic and
related structures such as the amygdala, orbitofrontal cortex, and hippocampus project to the ventral striatum (which includes
the nucleus accumbens),
which has connections through the ventral pallidum to the mediodorsal nucleus of the thalamus and thus to the prefrontal and cingulate cortices. |
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In one hypothesis, the striatum would be particularly in
involved in the selection of behavioral responses. |
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The selection of behavioral responses process may be achieved by a laterally spreading competitive interaction between striatal or pallidal neurons, which might be implemented by direct
connections between nearby
neurons in the striatum and globus pallidus. |
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Inhibitory interneurons within the striatum, via the interconnectivity of their dendrites, may play a part in the interaction
between striatal processing streams. |
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In addition to the response selection function by competition, the basal ganglia may enable
signals originating from non-motor parts of the cerebral cortex to be mapped onto motor signals to
produce behavioral output. |
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Short range integration of interactions within the
striatum may be
produced by the short length (for example 0.5 mm) of the intrastriatal axons of striatal neurons. |
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A strong
input to one part of
the striatum could spread like a
lateral competition signal. Such a mechanism could contribute to behavioral response selection in
the face of different competing input signals to the striatum. |
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Lateral inhibition could operate between striatal principle neurons by direct connections they receive from the cortex and could inhibit each other by their local axonal authorizations, which
spread in an area as large as their dendritic
trees, and which utilize GABA as their inhibitory transmitter. |
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Lateral inhibition could also operate in the pallidum and substantia nigra. Here again there are local axon collaterals as extensive
as the very large pallidal and nigral dendritic fields. |
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A selection
function between
processing streams in the basal ganglia, even without any
convergence anatomically between the processing streams, might provide an
important computational reason for the basal ganglia. |
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The direct inhibitory local connectivity
between the principal neurons within the striatum and globus pallidus would seem to provide a simple, and perhaps evolutionarily old, way in which to
implement competition between neurons in processing
streams. This might even be a primitive design
principle that characterizes the basal ganglia. |
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A system such as the basal ganglia with direct inhibitory recurrent collaterals may have evolved easily because it easier to make stable than architectures such is the cerebral
cortex with recurrent
excitatory connections. |
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The basal
ganglia architecture may have been especially
appropriate in motor systems in which instability could produce movement and coordination difficulties. |
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The hypothesis of lateral competition between
neurons of the basal ganglia can be sketched simply.
The inputs from
the cortex to striatum
are excitatory, and competition between striatal
neurons is
implemented by the use of an inhibitory
transmitter (GABA), and direct
connections between striatal neurons, within an
area which is approximately
coextensive with the dendritic arborization. |
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Given that the lateral connections between the striatal neurons are collaterals of the
output axons, the output must
be inhibitory onto the pallidal and nigral neurons. |
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To transmit
signals, neurons in the globus
pallidus and substantia
nigra must have high
spontaneous firing rates and respond by reducing their firing rates. |
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A simple, and perhaps evolutionary early, aspect of basal ganglia architecture is that
the striatal, pallidal,
and nigral neurons
implement competition
(for selection) by
direct inhibitory recurrent lateral connections. |
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Because of the slow conduction speed of dopaminergic neurons they are
probably only suitable for more tonic, long-term
adjustments of sensitivity. |
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One result of the convergence achieved by the medial pallidal/substantia nigra is
that even if inputs from different cortical
regions were kept segregated, there might nevertheless well be the possibility for a mutual competition between different pallidal
neurons, implemented by
interneurons. |
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Given the relatively
small number of neurons into which the cortical signals have been compressed, it would be feasible to
have competition implemented between the relatively small
population of neurons so that the competition would spread widely within these nuclei. This would allow selection by competition between these pathways, i.e. effectively
between information processing in different cortical areas. |
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Neural basis for 'habit' learning in which the basal ganglia have been implicated. |
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Storage of motor plans in the basal ganglia, which would be
instantiated as a series of lookups of the appropriate motor output
pattern to an evolving sequence of input
information. |
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Other parts of the motor system, such is a corticocortical pathways, may mediate the control of action in a voluntary, often slow way in the early stages of learning. |
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The corticocortical
pathways would set up the conditions, which
because of the continuing repetition, would be learned by the
basal ganglia. |
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If a strong
new pattern of inputs was received by the basal ganglia, this would result in a different
pattern of outputs being 'looked up' than that currently in
progress. |
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Outputs of
the globus pallidus
and substantia nigra
directed via the thalamus to motor regions such as a supplementary motor
cortex and the premotor
cortex potentially provide important output routes for the basal ganglia to produce actions. |
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There are also outputs of the basal ganglia to structures that may not be
primarily motor, such as a dorsolateral prefrontal cortex in
primates, and the inferior temporal visual cortex. |
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The hypothesis of basal ganglia function incorporates associative learning of cognitive inputs onto neurons, at both the cortico-striatal stages, and the striato-palladal and nigral stages. |
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Consistent with this hypothesis
of basal ganglia action it
has been possible to demonstrate long-term potentiation (LTP) in at
least some parts of
the basal ganglia. |
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Researchers have demonstrated LTP of limbic inputs to the nucleus
accumbens, and were able to show that such LTP is facilitated
by dopamine. |
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The ventral striatum would still be an
area allowing convergence of signals from the amygdala, orbitofrontal
cortex, and hippocampus. |
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A sparse distributed representation is a distributed representation in which
a small proportion of
the neurons is active
at any one time. |
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