Edmund
Rolls - Emotion Explained |
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Topic |
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Rolls; Emotion Explained |
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This book evolved from earlier
book The Brain and Emotion
(1999) |
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What produces emotions? -- reinforcing stimuli, i.e. rewards and punishers. |
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Why do we have emotions? -- emotions are evolutionarily adaptive as a provides an efficient way for genes to influence our behavior. |
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How do we have emotions? -- describing what is
known about brain mechanisms of emotion. |
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Why do emotional states feel like something? -- this is part of a
large problem of consciousness. |
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Functions of affective
states in motivated
behavior (including hunger,
thirst, and sexual behavior),. This book proposes a fundamental and simple
relation between emotion and motivation. |
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Interactions between mood, and cognition and memory. |
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Impulsive behavior that is a feature of borderline
personality disorder. |
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Emotional feelings, part of the much larger problem of consciousness. |
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Functional neuroimaging -- activations of some brain regions are directly correlated with subjective feelings of affective state. |
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What are emotions? Why do we have
emotions? What is their adaptive value? What are the brain mechanisms of emotion,
and how can disorders of emotion be understood? Why does it feel like something to have an
emotion? Why do emotions sometimes
feel so intense? |
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Emotion and
motivation are linked by the property that both involve rewards and punishers. |
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Emotions
can be thought of as states elicited by rewards or punishers. |
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Motivation
can be thought of as a state in which a reward is being sought, or a punisher is being avoided or escaped from. |
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Importance of reward and punishment for emotion and motivation. |
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Some stimuli
are innately rewarding or punishing and are called primary
reinforcers. |
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No learning
is necessary to respond to pain as aversive. -- Pain is a primary
reinforcer. |
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Stimulus-reinforcement
association learning. |
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Autonomic responses are those mediated through the autonomic
nervous system, via the vagus and sympathetic nerves, which affect smooth muscle. |
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Many endocrine
(hormonal) responses are mediated through the autonomic nervous system and are autonomic responses, for example the release of adrenaline
(epinephrine) from the adrenal
gland during emotional
excitement. |
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Previously neutral stimuli can by pairing with unconditioned stimuli come by learning the association to produce learned autonomic
responses. -- Classical
Conditioning |
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Classical conditioning is very similar to stimulus-reinforcer association
learning, except that in the case of classical conditioning the responses involved are autonomic and endocrine responses. |
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Motivation
refers to the state an animal is the and when it is willing to work for a reward or to escape from or avoid a punisher. |
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For both emotion and motivation, rewards and punishers are assessed in order to provide the goals for behavior. |
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Computing the reward and punisher value of sensory stimuli, and then using selection between different rewards and avoidance of punishers in a common reward-based currency appears to be the fundamental design that brains use in order to produce
appropriate behavior. |
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Having reward
and punishment systems is the solution that evolution has developed to produce appropriate
behavior. |
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Proposed definition of emotions -- emotions are states elicited by rewards and punishers. |
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Some Primary
Reinforcers (diagram) |
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A mood is a continuing state normally elicited by a reinforcer, and is thus part of what an
emotion is. |
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Functions of Emotion -- Reward,
Punishment, and Emotion |
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Rewards and Punishers |
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Stimulus-reward learning and reinforcement by rewards and punishers. |
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Selection of behavior -- cost-benefit 'analysis' |
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4 |
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Autonomic
and endocrine
responses |
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Emotional states are motivating |
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2 |
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63 |
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Brain Mechanisms Underlying
Emotion |
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Pathways
involved in emotion
(diagram) |
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Schematic diagram of connections of taste,
olfactory, somatosensory, and visual pathways. (diagram) |
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Taste |
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Smell |
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Pleasant and painful touch |
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Visual stimuli |
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Representing potential secondary
reinforces |
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Receptive field size and translation invariantce |
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An important question for
understanding brain function
is whether a particular object, (or face) is represented in the brain by the
firing of one or a few gnostic (or 'grandmother') cells, or whether instead
the firing of a group or ensemble of cells each with somewhat different
responsiveness provides the
representation. |
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Facial expression, gesture and view represented by a population of neurons in the cortex in the superior temporal sulcus. |
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The brain
mechanisms that build the appropriate view-invariant representations of objects required for learning emotional responses to objects, including faces. |
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Orbitofrontal Cortex |
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Phineas Gage |
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Effects of damage to the orbitofrontal cortex |
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Neurophysiology and functional neuroimaging of the orbitofrontal cortex |
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Taste |
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The nature of the representation of taste in the orbitofrontal cortex is that the reward value of the taste is represented. |
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Olfactory representation in the orbitofrontal cortex |
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Visual inputs
to the orbitofrontal cortex, and visual stimulus-reinforcer
association learning and reversal. |
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A representation
of faces in the orbitofrontal cortex |
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Cognitive influences on the orbitofrontal cortex |
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Human Orbitofrontal
Cortex |
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Executive function of the orbitofrontal cortex |
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The amygdala |
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Associative
processes involved in emotion-related learning |
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Pavlovian
or Classical Conditioning |
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Connections
of the amygdala |
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The amygdala is a subcortical region of 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 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 reinforcers, 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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Subcortical inputs to the amygdala include projections from the midline
thalamic nuclei, the subiculum, CA1
parts of the hippocampal formation, and the hypothalamus. |
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Emotions
are usually elicited by environmental stimuli analyzed to the object level, and not to retinal arrays. |
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Outputs of
the amygdala include
the well-known projections to the hypothalamus. |
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The amygdala has back projections to many areas of the temporal,
orbitofrontal, and insula cortices
from which it receives inputs. |
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There are separate output pathways from the amygdala for different fear related responses. |
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One output system of the amygdala is to the nucleus accumbens, a part of the striatum. |
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The core
part of the nucleus
accumbens is part of the pathway for approach responses to conditioned stimuli. |
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Consistently, dopamine release in the core part of the nucleus accumbens is increased by conditioned emotional stimuli, both appetitive and aversive. |
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Amygdala is
involved in responses made to stimuli that are associated by learning with primary reinforcers, including rewards as well as punishers. |
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Amygdala is
a brain region for stimulus-reinforcer
association learning. |
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The amygdala has partly dissociable
systems for Pavlovian
effects implemented via the central nucleus, and for effects of affective representations
implemented via the basolateral amygdala. |
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Neuronal activity in the primate
amygdala to reinforcing stimuli. |
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Responses
of amygdala neurons to
novel stimuli that
are reinforcing. |
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Neuronal responses in the amygdala to faces. |
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Cingulate Cortex |
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Mid-cingulate cortex, the cingulate motor area, and action-outcome learning. |
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187 |
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Human brain imaging investigations of mood and depression. |
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188 |
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Output Pathways for Emotional
Responses |
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Autonomic and Endocrine Systems |
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189 |
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Motor systems
for implicit responses,
including the basal ganglia. |
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Output systems for explicit responses to emotional stimuli. |
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Basal Forebrain and Hypothalamus |
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Basal Forebrain Cholinergic Neurons |
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Noradrenergic
Neurons |
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Effects of emotion on cognitive processing and memory. |
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195 |
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Whenever memories are stored, part of the context is stored with the memory. This is very likely to happen in associative neuronal networks such as those in the
hippocampus. |
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The CA3 part of the hippocampus may operate as a single associative memory capable of linking together almost arbitrary co-occurrences of inputs, including
inputs about emotional state that reach the entorhinal cortex from the amygdala. |
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Recall of a
memory occurs best in associative networks when the input key to the memory is nearest to the original input pattern of activity that was stored. |
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195 |
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Recall from
the hippocampus is likely to use the highly developed back
projections from the hippocampus to the neocortex. |
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195 |
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The hippocampus, which is implicated in in memory
of past episodes,
contains neurons that respond to combinations of spatial information and reward information. |
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196 |
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The ability to form associations between events including where they occur and what is present is the fundamental
property of episodic memory. |
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196 |
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The primate anterior hippocampus receives inputs from brain regions involved in reward processing such as the amygdala and orbitofrontal cortex. |
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Hippocampus
may store information about where emotion-related events happened; may take part in the recall of emotions when particular
places are seen again; and may provide a system in which the current mood can influence which memories are recalled. |
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Effects of mood on storage and recall could be facilitated by the back
projection from structures important in emotions such as the amygdala and orbitofrontal cortex to parts of the cerebral cortex important in representation of
objects,
such as the inferior temporal visual cortex. |
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Co-activity between forward inputs and back projections could result in facilitation or recall of cortical
representations (e.g. particular
faces)
that had become associated with emotional
states, represented by activity
in the amygdala. |
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198 |
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Theory of how effects
of mood on memory and perception could be implemented in the brain. Massive
projections from parts of the brain where mood is represented, such as the orbitofrontal cortex and amygdala, to the cortical
areas such is the inferior
temporal visual cortex and hippocampus-related areas
that project into these mood-representing areas. |
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Laterality effects in human emotional processing. |
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Hunger |
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Peripheral Signals for a Hunger and Satiety |
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224 |
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Control signals for a hunger and satiety |
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224 |
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Sensory-specific satiety |
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230 |
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Gastric distention |
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231 |
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Body fat regulation |
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232 |
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Conditioned appetite and satiety |
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233 |
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Brain Control of Eating and
Reward |
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233 |
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Hypothalamus |
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Neuronal activity in their lateral hypothalamus during feeding. |
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Hypothalamic neurons responsive to the sight, smell, and taste of food. |
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Effect of hunger |
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Sites in the hypothalamus and basal forebrain of neurons that respond to food. |
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Effects of signals related to hunger and satiety on hypothalamic neurons. |
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242 |
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Functions of the hypothalamus in feeding |
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243 |
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Brain mechanisms for the reward produced by the taste of food. |
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1 |
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Taste processing up to and including the primary
taste cortex of
primates is related to the identity of the tastant, and not to is reward value. |
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Taste and taste
related processing in the secondary taste cortex, including umami taste,
astringency, fat, viscosity, temperature and capsaicin. |
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The reward
value of taste is represented in the orbitofrontal cortex. |
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Convergence
between taste and olfactory processing to represent flavor. |
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254 |
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Brain mechanisms for the reward produced by the odor of food. |
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258 |
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Representation of information about odors by populations of neurons in the orbitofrontal cortex. |
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Responses
of orbitofrontal
cortex taste and olfactory neurons to the sight
of food. |
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Functions of the amygdala and temporal cortex in feeding. |
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Inferior Temporal Visual Cortex |
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Basal ganglia
are parts of the brain that include the striatum, globus
pallidus, substantia nigra, and subthalamic nucleus, and are necessary for the normal initiation of movement. |
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Depletion
of the dopaminergic
input to the striatum leads to the lack
in the initiation of voluntary
movement that occurs in Parkinson's
disease. |
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Basal ganglia
receive inputs
from all parts of the cerebral cortex, including the motor
cortex,
and have outputs
directed strongly toward the premotor and prefrontal cortex via which they could influence movement initiation. |
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The general connectivity of the basal ganglia is for cortical and limbic inputs to reach the striatum, which then project to the globus pallidus and substantia nigra pars reticulata, which in turn project via the thalamus back to the cerebral cortex. |
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Within the overall scheme of basal ganglia connectivity, there is a set of at least partially segregated parallel
processing streams. |
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There is evidence linking the ventral striatum and its dopamine input to reward. |
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Rats will self-administer amphetamines into the nucleus accumbens, and lesions of the nucleus accumbens attenuate the intravenous
self-administration of cocaine. |
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Lesions of
the dopamine pathways
that deplete the striatum of dopamine lead to a failure to orient to
stimuli,
a failure to initiate movements, catalepsy, and a failure to eat and drink. |
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325 |
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In humans, depletion
of dopamine in the striatum is found in Parkinson's disease, in which there is akinesia (i.e. a lack of voluntary movement), bradykinesia (slow movement), rigidity, and tremor. |
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The activity of many neurons in the putamen is related to movements. |
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There is a somatotopic organization of neurons in the putamen, with separate areas containing neurons responding to arm, leg, or orofacial movements. |
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The firing
rate of neurons in the putamen tends to be
linearly related to the amplitude of movements. |
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Patients
with basal ganglia disease frequently have difficulty in controlling the amplitude of their limb movements. |
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What computations are performed by the basal ganglia? |
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The striatum would be particularly involved in the selection of behavioral responses, and producing one coherent stream of behavioral output, with the possibility to switch to a different behavior if a higher priority input is received. |
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340 |
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The selection of behavioral responses may be achieved by laterally
spreading competitive interaction between striatal or palatal neurons. |
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Inhibitory interneurons within the striatum may play a part in the interaction between striatal processing streams. |
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A behavioral
selection function
between processing streams in the basal ganglia, even without any convergence anatomically between processing streams, might provide an
important computational rationale for the basal ganglia. |
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The basal
ganglia may be able to detect
combinations of conjunctively
active inputs
from quite widespread regions of the cerebral cortex using their combinatorial
architecture and a property of synaptic modifiability. In this way it would be possible to trigger any complex pattern of behavioral responses by any
complex pattern
of environmental inputs, using what is
effectively an associative network. |
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Dopamine
may play an important role in setting the thresholds of striatal neurons. |
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347 |
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Dopamine neurons could
not convey
information about a primary reward obtained if
the trial is successful, in the way that orbitofrontal cortex neurons do. |
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388 |
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Pheromones
can cause groups of women housed together to start cycling together. |
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400 |
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We may build
a computer that would perform
the functions of the emotions, yet we may not
want to ascribe emotional feelings to the computer. |
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We may think that the most important aspect of emotions is feelings and that their neural basis has not been accounted for. |
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How do the emotional
feelings and sensory
events come to feel
like anything? These 'feels' are called qualia.
They are great mysteries that have puzzled philosophers
for centuries. |
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Many actions can be performed
relatively automatically, without apparent conscious intervention, i.e. driving a car. |
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401 |
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Actions
that are relatively automatic could involve control of behavior by brain systems that are old in evolutionary
terms such in the basal
ganglia. |
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0 |
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401 |
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The basal
ganglia and cerebellum do not have backprojection
systems to most of the parts of the cerebral cortex from which they receive inputs. |
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0 |
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401 |
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Parts of the brain such as the hippocampus and amygdala involved in such
functions as episodic memory and emotion, do have major backprojection
systems to the high
parts of the cerebral cortex from which they receive forward projections. |
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401 |
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It may be that evolutionarily newer parts of the brain, such as the language areas and parts of the prefrontal
cortex, are involved in an alternative type of control of behavior, in which actions can be planned with the
use of a language system
that allows relatively arbitrary syntactic
manipulation of semantic
symbolic entities. |
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401 |
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The general view that there are many routes to behavioral output
is supported by the evidence that there are many
input systems to the basal
ganglia (from almost all areas of the cerebral cortex), and that neuronal activity in each part of the striatum reflects
the activity in the overlying cortical area. |
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The language areas offer one of many
routes to action, but a route particularly suited in planning actions, because of the syntactic manipulation of semantic entities, which may make
the long-term planning
possible. |
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General view of brain evolution and which, as areas of the cortex evolved, they
are laid on top of the existing circuitry connecting inputs to outputs, and in which each level in this hierarchy of
separate input and output data pathways may control
behavior are according to the specialized function it can perform. |
|
2 |
Rolls; Emotion Explained |
403 |
|
Mathematicians may get a hunch that something is
correct, yet not be
able to verbalize why. They may then resort to
formal, more serial
and language-like theorems to prove the case, and these seem to require conscious processing. |
|
0 |
Rolls; Emotion Explained |
427 |
|
The ventral
visual system is
concerned with selecting goals for action. It does this by providing invariant
representations of objects, with a representation
that is appropriate for interfacing to systems such as the amygdala and orbitofrontal cortex. |
|
24 |
Rolls; Emotion Explained |
427 |
|
The dorsal visual system helps with executing actions, for example in shaping the hand appropriately to pick up a selected object. Often
the sensori-motor operation is performed implicitly, i.e. without conscious awareness. |
|
0 |
Rolls; Emotion Explained |
427 |
|
Insofar as explicit
planning about future
goals and actions requires knowledge of objects and their reward or punisher associations, it is the ventral visual system that provides the appropriate
input for planning
future actions. |
|
0 |
Rolls; Emotion Explained |
427 |
|
When explicit, or conscious, planning is required,
activity in the ventral visual system will be closely
related to consciousness. |
|
0 |
Rolls; Emotion Explained |
427 |
|
It is to objects are represented in the ventral
visual system that we normally apply multistep planning processes. |
|
0 |
Rolls; Emotion Explained |
428 |
|
The outputs of the reward and punishment
systems must be treated by the action system as being the goals for action. |
|
1 |
Rolls; Emotion Explained |
428 |
|
The action
systems must be built to try to maximize the activation of the
representations produced by rewarding events, and to minimize the activation of the representations produced by punishers or stimuli associated with punishers. |
|
0 |
Rolls; Emotion Explained |
429 |
|
The architectural design
principles of the brain to the stage of the representation of rewards and punishers and thus of emotion seems apparent. |
|
1 |
Rolls; Emotion Explained |
429 |
|
How the brain
selects between reward and punishment signals how the cost of actions are taken into account, and how actions
are selected remain to be determined. |
|
0 |
Rolls; Emotion Explained |
454 |
|
Neural network simulation of biologically plausible pattern association memories such
as may be present in orbitofrontal cortex and amygdala and autoassociation or attractor networks. |
|
25 |
|
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