Edelman;
Wider than the Sky |
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Edelman;
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Some people
take the view that consciousness is a mere epiphenomenon with no material consequences. |
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We all know what consciousness is - it is what you lose when you fall into a deep
dreamless sleep and what you regain when you wake up. |
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Consciousness
is utterly dependent on the brain. |
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Consciousness emerges from the organization and operation of the brain. |
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When brain
function is curtailed, after strokes, and in certain limited phases of
sleep, consciousness is not present. |
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No return of the functions of the body and brain after death. Post mortem experience is simply not possible. |
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What features of the body and
brain are necessary and sufficient for consciousness to appear? |
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Consciousness
- "remembered present" - reflecting the fact that all past
experience is engaged in forming an integrated awareness of this single moment. |
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Primary consciousness is the state of being mentally
aware of things in the world, of having mental
images in the present. It is possessed by animals lacking semantic or
linguistic capabilities. |
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Higher-order consciousness involves the ability to be conscious
of being conscious. Recreate past episodes and form future intentions. |
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At a minimal level, higher-order consciousness
requires semantic ability, that is, the assignment of
meaning to a symbol. |
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In its most developed form, higher-order consciousness
requires linguistic
ability, that is, the mastery of a whole system
of symbols and a grammar. |
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Edelman;
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Different levels of consciousness - In REM sleep, dreams are conscious states. |
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In diseases
of consciousness, such as schizophrenia, hallucinations, delusions, and disorientations can occur. |
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In normal conscious states individuals experience qualia (e.g., greenness, warmth,
painfulness). |
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Qualia are high-order discriminations that
constitute consciousness. |
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Brain
contains at least 30 billion neurons. |
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Brain contains 1 million billion synapses. (1015) |
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About 200
different kinds of neurons in the brain. |
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Corpus callosum connects the two cortical hemispheres and consists of more
than 200 million reciprocal axons. |
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Hippocampus
is an evolutionarily ancient cortical structure, lined up like a pair
of curled sausages along the inner skirt of the temporal cortex. Cross section
looks like a sea horse. |
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Neural function of the hippocampus provides synaptic mechanisms underlying memory but should not be equated with memory itself.
While synaptic change is essential for the function of memory, memory is a system property of neural network connections. |
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Increased synaptic
strength or efficacy within a pathway leads to a higher
likelihood of conduction across a pathway. |
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A long-term
memory record results when particular synaptic connections between the hippocampus and the cortex are strengthened. |
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Basal ganglia
connect to the cortex via the thalamus. Reciprocal
connection patterns existing in the cortex itself
and between the cortex and the thalamus are lacking in the basal ganglia. |
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Activity of the basal ganglia is through inhibitory neurons using GABA has neurotransmitter. Since inhibition
of inhibition (or disinhibition) can occur in these loops, they can stimulate
target neurons as well as suppress their
activity. |
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Edelman;
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Basal ganglia
are believed to be involved in the initiation and
control of motor
patterns. |
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Much of procedural memory (e.g., ride a
bicycle) and other nonconscious learned activity depends on the functions of the basal
ganglia. |
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Edelman;
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Value systems,
by projecting profusely,
affect large populations of neurons simultaneously. |
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Value system
bias neuronal responses
affecting both learning and memory necessary for survival. |
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Three main neuroanatomical motifs in the
brain: (1) thalamocortical, (2) polysynaptic loop structure, (3) diffuse ascending projections of
the different value systems. |
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No two individuals, not even identical twins, possess the same anatomical pattern. |
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Earliest stages of development, the cellular organization characteristic of a species, is
controlled by families of genes, including the Hox
genes. |
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At a certain
point in development, the control of neural connectivity
becomes epigenetic;
it is not prespecified
as "hardwiring" by genes,
but rather is guided by
patterns of neural activity. Neurons that "fire together,
wire together." |
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Result is a pattern
of constancy and variation leading to highly individual networks in each
animal. |
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Ablation of cortical area V1
leads to blindness. |
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There are areas of the brain
that if damaged or removed will lead to permanent unconsciousness:
midbrain reticular formation; region of the thalamus containing the intralaminar nuclei. |
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Consciousness
is comprised of a dynamic interaction of multiple brain regions. |
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One simple principle governs how
the brain works: it evolved; It was not designed. Natural selection acting in complex environments over eons of time. |
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Neural Darwinism - the theory of neuronal group
selection. |
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Explanation of consciousness will require an
understanding of perception, memory, action,
attention. |
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Extraordinary amount of variation in each individual brain. |
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Enormous variation at the levels of neuronal
chemistry, network structure, synaptic strength, temporal properties,
memories, and
motivational patterns governed by value systems. |
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Edelman;
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In sensory
systems such as vision, multiple cortical regions that are functionally segregated (e.g., color, movement, orientation), can exceed thirty in number, distributed all over
the brain. |
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No superordinate area or executive program binding these functions together
into a coherent percept. |
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Remarkable tendency of brains to
seek closure and avoid gaps. |
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Closure phenomena - responses to strokes, extraordinary adaptation and integration, damaged brain responds to the loss of cortical area. |
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Brains of
higher-level animals autonomously construct patterned responses to environments that are full of novelty. |
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Edelman;
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Theory of Neuronal Group Selection, TNGS -
three tenets: (1) Developmental selection, (2) Experimental selection, (3)
Reentry |
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Developmental selection - during early development,
epigenetic variations in the patterns of
connections, create repertoires in each brain area; millions of variant
circuits or neuronal groups. Neurons that "fire together wire together." |
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Experimental selection - large variations in synaptic strength, positive and
negative, result from variations in environmental
input during behavior. Synaptic modifications are
subject to the constraints of value systems. |
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Reentry -
during development, large numbers of reciprocal connections are
established both locally
and over long distances.
This provides a basis for signaling between mapped
areas across reciprocal fibers. |
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Reentry - recursive interchange of parallel signals among brain areas, coordinate the activities of
different brain areas
in space and time. |
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Unlike feedback, reentry is not a sequential transmission of an error signal in a simple loop.
Instead, it simultaneously involves many parallel
reciprocal paths and has no prescribed error
function. |
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Consequence
of this dynamic process
is the widespread synchronization of the activity of widely
distributed neuronal groups. Binds their functionally segregated activities
into coherent output. |
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Reentry is
the central organizing principle that governs the spatiotemporal
coordination among multiple
selectional networks of the brain. |
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Reentry
solves the binding problem. Through reentry, for example, the color, orientation and
movement of a visual object can be integrated. |
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Various individual
maps that are functionally
segregated can be coordinated by communicating directly with each other via reentry. |
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It is the neuronal
groups made up of excitatory
and inhibitory neurons in particular anatomical patterns,
rather than individual neurons, that are selected. |
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Proposal of the TNGS in 1978, neuronal groups connected by reentrant interactions are the selectional units in higher-level
brains. |
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Edelman;
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Brain is so
versatile in its responses because those responses are degenerate. Degeneracy is the ability of structurally
different elements of a system to perform the same function. |
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Edelman;
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Genetic code
-- Code words are degenerate. Third position of many triplet codons can contain any one of the four letters
without changing their coding specificity. |
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Identical twins who have similar immune responses to a foreign agent do not generally use identical combinations of antibodies
to react to that agent. |
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Edelman;
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Many structurally
different antibodies with similar specificities can be selected in the immune response to
a given foreign molecule. |
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Edelman;
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Mutual reentrant interactions, for a brief time, link various
neuronal groups in each map to those of others to
form a functioning circuit. Neurons that yield such circuits fire more or less in phase with each other, or synchronously. |
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In the next time period, different neurons and neuronal groups may form a structurally different
circuit, which nevertheless has the same output. |
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These different
circuits are degenerate - they are different in structure, but they yield similar outputs to solve the binding problem. |
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Edelman;
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As a result of
reentry, the properties of synchrony and coherency allow more than one structure to give a similar output. |
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Edelman;
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As long as such degenerate operations occur in succession to link distributed populations of neuronal groups, there is no need for an executive. |
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Edelman;
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Different cell can carry out the same function, and the same cell can, at two different times, carry out different functions in different neuronal groups. |
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Edelman;
Wider than the Sky |
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In the mammalian nervous system,
perceptual categorization is
carried out by interactions between sensory and motor systems by what Edelman calls global mapping. |
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Edelman;
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Global mapping is a dynamic
structure containing various
sensory maps, each with different functionally
segregated properties, linked by reentry. |
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Edelman;
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The global
mapping is linked by non-reentrant connections to motor maps and subcortical systems such as the cerebellum and basal ganglia. |
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Edelman; Wider than the Sky |
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Higher-order cortical maps in the prefrontal, parietal, and temporal areas carry out
this global mapping.
No linear sum of global mappings could give rise to such a generalization.
Instead, the generalization arises by abstracting
certain features of
the mappings by higher-order maps. |
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Edelman;
Wider than the Sky |
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Perceptual categorization and concept formation would not be adaptive to an animal in the absence of memory. |
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Edelman;
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Understanding of memory is essential for
formulating a theory of consciousness. |
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Edelman;
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Memory
results from changes in synaptic efficacy
(synaptic strength) in circuits of neuronal groups. |
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Edelman;
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After changes have occurred,
they tend to favor recruitment
of certain of these circuits to yield reenactment. This recruitment may occur in
more than one way - it may be degenerate. |
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Edelman;
Wider than the Sky |
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Certain forms of memory require fast changes in synaptic efficacy in time periods
less than about one-third of a second. |
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Edelman;
Wider than the Sky |
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Other forms of memory require slower, but more stable, changes in synaptic strength. |
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Edelman;
Wider than the Sky |
51 |
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Episodic memory depends on interactions between the hippocampus and the cerebral cortex. |
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Edelman;
Wider than the Sky |
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Procedural memory - reflects motor learning |
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Edelman;
Wider than the Sky |
51 |
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Episodic memory - long term recall of sequences of events or
narratives. Depends on interactions between the hippocampus and the cerebral cortex. |
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Edelman;
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Memory cannot simply be equated
with synaptic change, although changes in synaptic strength are essential for
it. |
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Edelman; Wider than the Sky |
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Each event of memory is a
dynamic and context-sensitive. Memory involves a repetition of a mental
act that is similar
to but not identical with previous acts. Memory is recategorical; it does
not replicate an original
experience exactly. |
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Edelman;
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Memory
should be looked on as a property of degenerate
nonlinear interactions in a multidimensional network of neuronal groups. Such interactions
allow a non-identical "reliving" of a set of prior acts and events. |
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Edelman;
Wider than the Sky |
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Memories
are necessarily associative and never identical. |
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Edelman;
Wider than the Sky |
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Mechanisms leading to perceptual categorization - global
mappings, concept formation, dynamic short-term memory - all call upon three
major motifs. |
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Edelman;
Wider than the Sky |
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(1) Thalamocortical maps |
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Edelman;
Wider than the Sky |
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(2) Subcortical organs concerned
with temporal succession
(hippocampus, basal ganglia, cerebellum) |
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Edelman;
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(3) Diffuse
ascending value systems. |
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Edelman;
Wider than the Sky |
53 |
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Value-category memory system - constraints of value
systems can determine the degree and extent of recall. |
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Edelman;
Wider than the Sky |
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At a point in evolutionary
time corresponding to the transition between reptiles and birds and between reptiles and mammals, a new reciprocal connectivity appeared
in the thalamocortical system. |
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Edelman;
Wider than the Sky |
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Massively reentrant connectivity developed
between the cortical areas carrying out perceptual
categorization and the more frontal areas responsible for
value-category memory based on fast changes in
synaptic strength. |
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Edelman;
Wider than the Sky |
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Dynamic reentrant interactions in the thalamocortical system. |
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Edelman;
Wider than the Sky |
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Bootstrapping between memory and perception is assumed to be stabilized within time periods ranging from hundreds
of milliseconds to
seconds. |
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Edelman;
Wider than the Sky |
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"Remembered Present" -- dynamic interaction between memory and ongoing perception that gives rise to consciousness. |
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Edelman;
Wider than the Sky |
55 |
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Activity of the reticular nucleus of the thalamus gates various combinations of activity of those specific thalamic nuclei
corresponding to different sensory modalities. |
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Edelman;
Wider than the Sky |
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Intralaminar nuclei of the
thalamus, which send diffuse
connections to most
areas of the cortex, helps to synchronize thalamocortical responses and regulate the overall levels of
activity in these multiple reentrant systems. |
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Edelman;
Wider than the Sky |
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Reentrant pathways leading to primary consciousness - (diagram) |
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Edelman;
Wider than the Sky |
57 |
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Ability to create a scene by reentrant correlations between value-category memory and perceptual categories is the basis for the emergence of primary
memory. |
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Edelman;
Wider than the Sky |
57 |
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Some of the earliest
categorizations are related to signals from the animal's own body and brain. |
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Edelman;
Wider than the Sky |
57 |
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Signals from "self"
systems begin even before birth and remain a
central feature of primary consciousness. |
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Edelman;
Wider than the Sky |
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Memories
conditioned by the history of reward and punishment during an animal's past behavior
- key role in emotional responses and associated feelings. |
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Edelman;
Wider than the Sky |
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Ability to construct a conscious scene in a fraction of a second is the ability
to construct a remembered
present. |
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Edelman;
Wider than the Sky |
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Higher-order consciousness confers the ability to imagine
the future, explicitly recall
the past, and to be conscious
of being conscious. |
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Edelman;
Wider than the Sky |
67 |
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Brain complexity via differentiation and
integration - contrast with (1) perfect gas, which had no integration and (2) perfect crystal, which has
integration but no variety. - (diagram) |
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Edelman;
Wider than the Sky |
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Responses triggered by the reentrant dynamic core can also
stimulate non-conscious responses through the basal ganglia network. These travel along parallel, polysynaptic, one-directional pathways that leave the cortex, reach the various components of the
basal ganglia and certain thalamic nuclei, and finally return to the cortex. |
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Edelman;
Wider than the Sky |
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Responses subserving consciousness can connect to activity patterns in unconscious areas, served mainly but not exclusively by the basal ganglia. |
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Edelman;
Wider than the Sky |
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Dynamic Core
- (diagram) |
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Edelman;
Wider than the Sky |
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Basal ganglia
consist of the striatum (caudate nucleus and putamen), globus
pallidus, substantia nigra, and subthalamic nucleus. |
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Edelman;
Wider than the Sky |
89 |
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A major portion of the basal ganglia, constituting input
nuclei from the cortex, is the striatum, which consists of the caudate nucleus and putamen. |
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Edelman;
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89 |
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Globus pallidus and one part of the substantia
nigra make up the major output nuclei projecting to the thalamus. |
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Edelman;
Wider than the Sky |
89 |
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Basal ganglia
receive inputs from practically all regions of
the cortex. |
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Edelman;
Wider than the Sky |
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Basal ganglia
project to the prefrontal
cortex and to the association
areas, which weigh decisions
related to action. |
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Edelman;
Wider than the Sky |
91 |
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Motor circuit of the Basal
Ganglia - (diagram) |
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Edelman;
Wider than the Sky |
91 |
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Connections between basal ganglia and cortex are involved in the
execution of automatic
motor programs.
[FAPs] |
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Edelman;
Wider than the Sky |
91 |
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During conscious
learning of tasks, a considerable amount of cerebral cortex is engaged. |
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Edelman;
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With practice, conscious attention is not required, and acts become automatic. |
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Edelman;
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An attractive hypothesis is that
collaborations between the cortex and basal ganglia set up synaptic changes that lie behind procedural learning. |
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Edelman;
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93 |
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For example, practicing
musical passages will eventually result in the
ability to "rattle them off" without detailed attention. |
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Edelman;
Wider than the Sky |
94 |
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A pianist
playing a concerto may execute
passages without conscious attention, note by
note, but simultaneously may plan consciously or think
ahead about an upcoming musical
phrase or tempo. |
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Edelman;
Wider than the Sky |
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Certain portions of the cortex may be engaged in interactions with the basal ganglia
without being directly involved in the operation of the dynamic
core. |
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Edelman;
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Evolution of higher-order
consciousness. (diagram) |
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Wider than the Sky |
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