Rolls
& Deco; Noisy Brain |
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& Deco; Noisy Brain |
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The relatively random spiking times of individual neurons produces a source of noise in the brain. |
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& Deco; Noisy Brain |
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Functional neuroimaging is useful to indicate where in the
human brain different processes take place, and
the show which functions
can be disassociated from each other. |
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There are often 5000-20,000 inputs per neuron. |
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& Deco; Noisy Brain |
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A learning rule presaged by Donald Hebb (1949) proposes that synapses increase in strength when
there is conjunctive presynaptic and postsynaptic activity. |
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& Deco; Noisy Brain |
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Long-term potentiation is a use-dependent and sustained increase in synaptic
strength that can be induced by a brief periods of synaptic stimulation. |
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LTP is long-lasting, develops rapidly, is synapse specific, and in some cases
is associative. It is of interest as a potential synaptic mechanisms underlying some forms of memory. |
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Systems-level on Analysis of
Brain Function |
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& Deco; Noisy Brain |
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Pathways involved in some
sensory systems (diagram) |
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Find structure and connectivity
of the neocortex |
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Excitatory cells in connections |
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Inhibitory cells and connections |
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& Deco; Noisy Brain |
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Basket cells
comprise 5-7% of the
total cortical cell population, compared with approximately 72% of pyramidal
cells. |
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& Deco; Noisy Brain |
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Basket cells
receive synapses from the main extrinsic afferents to the neocortex, including thalamic afferents, so that they must
contribute to a feedforward inhibition of pyramidal cells. |
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Typical quantitative estimates
for neocortex (table) |
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& Deco; Noisy Brain |
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Local recurrent connectivity implementing local reentrantcy is a feature of cortical
computation. |
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The number of recurrent connections per neuron
is in the order of 10,000. |
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A principle
of cortical design is that it does have in part local connectivity, so that each part can have its own processing and
storage, which may be triggered
by other modules, but is a distinct operation from that which occurs
simultaneously in other
modules. |
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& Deco; Noisy Brain |
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Functional pathways through cortical layers. |
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In primary
sensory cortical areas, the main extrinsic forward input is
from the thalamus, and ends in layer 4, where synapses are formed onto spiny
stellate cells.
These in turn project heavily onto pyramidal cells in layers 3 and 2, which in turn send projections
forward to the next
cortical area. |
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The circuitry is more complex
because the thalamic afferents
also synapse onto the basal dendrites in or close to the layer 2
pyramidal cells, as well is onto layer 6
pyramidal cells and inhibitory interneurons. |
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Forward corto-cortical afferents to a cortical area sometimes have a columnar pattern to their
distribution, with a column width 200-300 µ in diameter. |
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& Deco; Noisy Brain |
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Theoretical Significance of Back Projections in the Neocortex |
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& Deco; Noisy Brain |
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Cortical processing takes place through a hierarchy of cortical stages. Convergence and competition are key aspects of the processing. |
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& Deco; Noisy Brain |
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Neurons in
each cortical stage respond for 20-30 ms when an object can just be seen. |
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& Deco; Noisy Brain |
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The time for processing to travel from V1 to the inferior temporal visual cortex (IT) is approximately 50 ms. |
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Backprojections are a major feature of cortical connectivity. |
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& Deco; Noisy Brain |
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Some of the functions of back
projections are in memory recall, in forming attractor networks in which there
are associatively modifiable synaptic connections in both the feedforward and feedback connections between adjacent
levels in the hierarchy, and in top-down attentional modulation implemented by biased competition. |
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Orbitofrontal cortex, amygdala and hippocampus are stages of information processing at which the different sensory modalities (such
as vision, hearing, touch, taste, and smell for the orbitofrontal cortex and
amygdala) are brought together, so that correlations between
inputs in different
modalities can be detected
in these regions, but not
at prior cortical processing stages in each modality, as these cortical processing stages are mainly unimodal. |
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& Deco; Noisy Brain |
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As a result of bringing together any two sensory modalities, significant correspondences between the two modalities can be detected. |
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& Deco; Noisy Brain |
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A highly developed system of excitatory recurrent collateral
connections between nearby
pyramidal cells is a hallmark of neocortical design. |
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Autoassociative memories, or attractor neural networks, store memories, each one of which is represented by a pattern of neural activity. |
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& Deco; Noisy Brain |
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Memories
are stored in the recurrent synaptic connections between the neurons of the network, for example in the recurrent collateral connections
between cortical pyramidal cells. |
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& Deco; Noisy Brain |
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Autoassociative networks can recall
the appropriate memory from the network when provided with a fragment of one
of the memories. This is called completion. |
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& Deco; Noisy Brain |
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A feature of autoassociative
memory is that it is content
addressable; i.e. the information
in the memory can be accessed if just the contents of the memory (or in part of the
contents of the memory) are used. |
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Autoassociation memory can be used as a short-term memory, in which iterative processing round the recurrent collateral
connection loop keeps a representation
active by continuing
neuronal firing. |
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The short-term
memory reflected in continuing neuronal firing for several hundred milliseconds after
a visual stimulus is removed, which is present in visual
cortical areas such as inferior
temporal visual cortex, is probably implemented
in this way. |
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Autoassociation memory appears to be used in a short-term
memory in the prefrontal
cortex. Temporal visual cortical areas have
connections to the ventrolateral prefrontal cortex that help to implement short-term
memory for visual
stimuli. |
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& Deco; Noisy Brain |
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Parietal cortex has connections to dorsolateral prefrontal cortex for
the short-term memory
of spatial responses. |
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& Deco; Noisy Brain |
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Prefrontal short-term
memories provide a mechanism that enables attention to be maintained through
backprojections from
the frontal cortex areas
to the temporal and parietal areas that send connections to the prefrontal cortex. |
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& Deco; Noisy Brain |
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Attractor networks are implemented by the forward and backward connections between cortical areas. |
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& Deco; Noisy Brain |
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An autoassociation network with the rapid synaptic plasticity can learn each memory in one trial. Because of its one-shot rapid learning, and
ability to complete,
this type of network is well suited for episodic
memory storage, in which each past episode must be stored and recalled later from fragments, and kept separate from other episodic memories. |
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& Deco; Noisy Brain |
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Perhaps the most important and
useful property of autoassociative memories is that they complete an incomplete input vector, allowing recall of a whole
memory from a small
fraction of it. |
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& Deco; Noisy Brain |
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The memory
recalled in response to a fragment is that stored in the
memory that is closest
in pattern similarity (as measured by the dot product, or correlation). |
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& Deco; Noisy Brain |
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Because the recall in associative memory is iterative and progressive, the recall can be perfect. |
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& Deco; Noisy Brain |
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If the network is modeled as 'integrate-and-fire' neurons,
it can effectively relax
into its recall state very rapidly, in one or two time constants of the synapses. This corresponds to perhaps 20 ms in the brain. |
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& Deco; Noisy Brain |
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One factor in the rapid dynamics of associative networks with
brain-like 'integrate-and-fire' membrane and synaptic properties is that with ongoing spontaneous activity, some of the neurons in the network are close to threshold already, before the recall cue is applied,
and hence some of the neurons are very quickly
pushed by the recall cue
into firing, perhaps
within 1-2 ms. |
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& Deco; Noisy Brain |
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In a sparse
distributed representation, a small proportion of the neurons its active at any one time to represent any one stimulus or event. |
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& Deco; Noisy Brain |
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One advantage of distributed encoding is that the similarity between two
representations can be reflected by the correlation between the two patterns of activity that
represent the different stimuli. |
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& Deco; Noisy Brain |
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Another advantage of distributed encoding is that the number of different stimuli
that can be represented
can be very large. |
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& Deco; Noisy Brain |
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Information can be read from the ensemble of neurons using a simple measure of the similarity
of vectors, the correlation (or dot product). |
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& Deco; Noisy Brain |
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Vector similarity operations characterize the operation of many neuronal networks. |
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& Deco; Noisy Brain |
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Neurophysiological results show that both the ability to reflect
similarity by vector
correlation, and the utilization of exponential coding capacity, are
properties of real neuronal networks in the brain. |
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The brain
codes stimuli in such a way that the code can be read off a simple dot product or correlation-related decoding. |
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The spiking of typical cortical neurons is close to that of a Poisson
process, i.e. for a given mean firing rate, the emission times of the spikes are approximately random and
independent of each
other. |
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& Deco; Noisy Brain |
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The time
constants of the synapses are an important factor in determining how noise, including that from the stochastic firing of other neurons, affects the operation of the network. |
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& Deco; Noisy Brain |
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Whereas the AMPA and GABA receptors operate with relatively short
time constants (in the
order of 5 ms), the NMDA receptors have a much longer time constant (in the order of 100 ms), which dampens the effect of a high frequency
noise and helps to promote
stability and dampen oscillations in the cortical network. |
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& Deco; Noisy Brain |
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The neuronal
membrane and dendritic
time constants are in
the order of 20 ms. Their effect on
the temporal dynamics of the network is less than that of the synapses. |
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& Deco; Noisy Brain |
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The overall
temporal properties of the network will be influenced by the synaptic and membrane time constants. |
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& Deco; Noisy Brain |
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Given that the
time constants of GABA
synapses is in the order of 10-20 ms, of AMPA receptors is 2-5 ms, and NMDA receptors is approximately 100 ms, oscillations are less likely to occur when the system is
strongly activated so that the neurons are firing
fast, and the voltage-independent NMDA receptors are contributing significantly to the currents in the system. |
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& Deco; Noisy Brain |
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Information theory provides the means for quantifying how much neurons communicate to other neurons, and thus provide a
quantitative approach to fundamental questions about information processing in the brain. |
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& Deco; Noisy Brain |
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We must investigate to what
extent the information provided by different cells is redundant or instead independent. |
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& Deco; Noisy Brain |
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Response of single
neurons in the inferior
temporal visual cortex to sets of objects and/or faces. |
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& Deco; Noisy Brain |
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Across a population
of cells, the firing rate information provided by each neuron tends to be independent; i.e. the information increases approximately
linearly with the number of neurons. |
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& Deco; Noisy Brain |
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Linear increase in information with the number of neurons has been found in many research
studies, including for face and object representations in the inferior temporal visual cortex,
for neurons responding to spatial view in the hippocampus, for neurons responding to head
direction in the presubiculum, and for neurons responding to olfactory and taste stimuli in the orbitofrontal cortex. |
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& Deco; Noisy Brain |
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The number
of stimuli that can be
encoded rises exponentially as the number of neurons in the ensemble increases. |
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& Deco; Noisy Brain |
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The information in the firing rate across a population of neurons can be read
moderately efficiently by decoding procedure as simple as a dot product. |
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& Deco; Noisy Brain |
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The dot
product operation is the simplest type of processing that
might be performed by neuron, as it involves taking a dot
product of the incoming firing rates with the receiving synaptic
weights to obtain the activation
of the neuron. |
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& Deco; Noisy Brain |
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Dot product encoding ensures that the simple emergent properties of associative neuronal networks such
as generalization, completion, and graceful degradation can be realized very naturally and simply. |
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& Deco; Noisy Brain |
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There is little
additional information, to the great deal available in the firing rates, from any stimulus-dependent cross-correlations of synchronization that may be present. |
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& Deco; Noisy Brain |
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Stimulus dependent synchronization might only be useful for grouping different neuronal populations, and would not easily provide a
solution to the binding
problem in vision. |
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& Deco; Noisy Brain |
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The binding
problem in vision may be solved by the presence of neurons that respond to combinations of features in a given spatial position with respect to each other. |
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& Deco; Noisy Brain |
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There is little
additional information to the great deal
available in the firing rates, from the order on which spikes
arrive from different
neurons. |
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& Deco; Noisy Brain |
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The spiking
of neurons in the brain is almost random in time for a given mean rate, i.e. the spiking is approximately Poisson, and
this randomness
introduces noise into
the system, which makes the system behave stochastically. |
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& Deco; Noisy Brain |
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The integrate-and-fire approach is a computationally realizable way to simulate the operation of neural networks when operating stochastically. |
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& Deco; Noisy Brain |
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In the simulation, each of the neurons show spiking that it's very much like that of neurons recorded in the
brain. |
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& Deco; Noisy Brain |
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Autoassociation attractor
systems have two types of stable fixed points: (1) a spontaneous state with a low firing rate, and (2) one or more attractor
states in which the positive
feedback implemented by the recurrent collateral connections
maintain a high firing rate. |
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The high
firing rate attractor state is sometimes referred
to as the persistent state, because the high firing rate
persists to maintain a
set of neurons active, which might implement a short-term memory. |
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& Deco; Noisy Brain |
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The stable
points in the system can be visualized in an energy landscape. |
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& Deco; Noisy Brain |
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The area in the energy landscape within which the system will move to a stable attractor state is call the basin of attraction. |
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& Deco; Noisy Brain |
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The attractor dynamics can be pictured by energy
landscapes, which indicate the basins of attraction by valleys, and the attractor states or fixed points
by the bottom of the valleys. |
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The stability
of an attractor is characterized by the average time in which the system stays in the basin of attraction under the influence of noise. |
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Noise
provokes transitions
to other attractor states. |
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& Deco; Noisy Brain |
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Two factors determined
the stability: (1) the depths of the attractors, and (2) high noise will make it more likely that the system will jump over an energy boundary from one
state to another. |
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The noise could arise not only from the probabilistic
spiking of the neurons, but also from any other source of
noise in the brain or environment, including the effects that is distracting
stimuli. |
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Neurons are
held close to (just slightly below) their firing threshold so that any incoming input can rapidly cause sufficient further depolarization to produce a spike. |
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It is this ability to respond rapidly to an input, rather
than than having to charge up the cell membrane from the resting potential to the threshold, that enables neuronal networks in the brain, including attractor
networks, to operate
and retrieve information so rapidly. |
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Charging up
to cell membrane from
the resting potential
to the threshold is a slow process determined by the time constant of the neuron and
influenced by the synapses. |
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Spike trains
are essentially Poisson-like because the cell potential hovers
noisily close to the threshold
for firing, the noise being generated in part by the Poisson-like
firing of the other
neurons in the network. |
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& Deco; Noisy Brain |
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Noise and spontaneous firing help to ensure
that when a stimulus arrives, there are always some neurons
very close to threshold that respond rapidly, and then communicate their firing to other neurons through the modified synaptic weights, so that
an attractor process
can take place very rapidly. |
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& Deco; Noisy Brain |
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Noise sources
in the brain arise from effects that make firing
properties of neurons very close to Poisson. |
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If the system has a finite number of neurons, then the Poisson or semi-random effects will not average out, and the average firing rate of the population will fluctuate statistically. |
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Signal-to-noise ratio can be measured in a network by the average
firing rate change produced by the signal in the neuronal population
divided by the standard deviation of the firing
rates. |
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& Deco; Noisy Brain |
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Noise inherent in brain activity has a number of advantages by making the dynamics stochastic, which allows for many remarkable
features of the brain, including creativity, probabilistic decision
making, stochastic
resonance, unpredictability, conflict resolution,
symmetry breaking, allocation to discrete categories, and many of the important properties. |
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& Deco; Noisy Brain |
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Statistical fluctuations are not just a concomitant of
neural processing, but can play an important and unique role in cortical function. |
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& Deco; Noisy Brain |
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Computations
can be performed through stochastic dynamical
effects, including the role of noise in enabling probabilistic jumping across the barrier in the energy landscape describing the flow of the
dynamics in attractor
networks. |
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& Deco; Noisy Brain |
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Stochastic dynamical effects help to understand short-term
memory, attention, long-term memory, and decision-making in the brain, and
to establish a link between neuronal firing
variability in probabilistic
behavior. |
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Mean-field based theory -- starting from the mathematical models of biologically realistic single neurons
(i.e. spiking neurons),
one can derive models that describe the joint activity of pools (or populations) of equivalent neurons. The computational unit is a neuronal pool, population, or assembly. |
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Short-Term Memory and Stochastic
Dynamics |
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& Deco; Noisy Brain |
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Cortical short-term memory systems and attractor networks |
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& Deco; Noisy Brain |
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There are a
number of different short-term
memory systems, each implemented in a different cortical area. |
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The short-term
memories may operate as autoassociative
attractor networks. |
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The short-term
memory is implemented by subpopulations of neurons that
show maintained activity in a delay., while a stimulus or
event is being remembered. |
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The autoassociation could be implemented by associatively modifiable
synapses between connected
pyramidal cells within an area, or by the forward and backward connections
between adjacent cortical areas in a hierarchy. |
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One short-term
memory system is in the dorsolateral
prefrontal cortex, area 46. This is involved in remembering the locations of spatial responses. |
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There is another short-term memory system in a more
dorsal and posterior part of the prefrontal cortex involved in remembering the
position in visual
space to which an eye
movement (a saccade) should be made. In this case the short-term
memory function is topographically
organized. |
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Local cortical connections between nearby pyramidal cells implement an attractor network. |
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& Deco; Noisy Brain |
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Another short-term
memory system is implemented in the inferior temporal visual cortex. |
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Some neurons respond more to a novel than a familiar visual stimulus. |
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& Deco; Noisy Brain |
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A possible cortical
model of a topographically
organized set of attractor
networks in the prefrontal
cortex that could be used to remember the
positions to which saccades should be made. (Diagram) |
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Short-term memory is also found in the inferior
frontal convexity cortex in a region connected to the ventral temporal cortex. |
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Prefrontal cortex short-term memory networks, and
their relation to temporal and parietal perceptual networks. |
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& Deco; Noisy Brain |
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A common way that the brain uses
to implement a short-term
memory is to maintain the firing of neurons during a short memory period after the end of a stimulus. |
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In the inferior temporal cortex the firing of neurons for short-term memory may be maintained for a few hundred milliseconds. In more ventral temporal cortical
areas such as the entorhinal
cortex the firing may
be maintained for longer
periods.
In the prefrontal
cortex the firing may
be maintained even for tens
of seconds. |
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& Deco; Noisy Brain |
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In the dorsolateral and inferior convexity prefrontal
cortex the firing of
the neurons may be related
to the memory of spatial
responses or objects. |
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& Deco; Noisy Brain |
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The firing
of neurons may be maintained by the operation of associatively
modified recurrent collateral connections between nearby pyramidal cells producing attractor states in autoassociative
networks. |
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& Deco; Noisy Brain |
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For short-term
memory to the maintained during periods in which new
stimuli are to be perceived, and must be separate networks for perceptual and short-term memory functions. |
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& Deco; Noisy Brain |
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Two coupled networks, one in the inferior temporal visual cortex for
perceptual functions,
and another in the prefrontal cortex for maintaining the short-term memory during intervening stimuli, provide a precise model of the interaction of perceptual and short-term memory systems. |
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& Deco; Noisy Brain |
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A prefrontal
cortex attractor (autoassociation) network could
be triggered by a sample visual stimulus represented
in the inferior temporal visual cortex and keep this attractor active during a memory interval in which intervening stimuli are shown. |
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& Deco; Noisy Brain |
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For ongoing perception to occur unhindered, implemented by posterior cortex (parietal and temporal lobe) networks, there must be a separate
set of modules that is capable of maintaining a representation over intervening stimuli. |
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A short-term
memory autoassociation network in the prefrontal cortex could hold active a working memory representation by maintaining its firing in an attractor state. (diagram) |
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95 |
|
The ability of the dorsolateral prefrontal cortex to provide multiple separate short-term attractor
memories could provide the basis for its function
in planning. |
|
0 |
Rolls
& Deco; Noisy Brain |
95 |
|
Neuronal activity in the inferior temporal visual
cortex (IT) is driven by each new incoming visual stimulus,
whereas in the prefrontal cortex, neurons start to fire when the sample stimulus is shown, and continue the firing that represents the sample stimulus, even
when potential match stimuli are being shown. |
|
0 |
Rolls
& Deco; Noisy Brain |
95 |
|
Computer simulation shows that a crucial parameter is the strength of the intermodular connections between the prefrontal cortex and the posterior cortex (temporal or parietal cortex). |
|
0 |
Rolls
& Deco; Noisy Brain |
97 |
|
Computer simulation shows that the operation of the prefrontal
cortex in short-term
memory and its relation to posterior perceptual networks, can
be understood by the interaction of two
weakly coupled attractor networks. |
|
2 |
Rolls
& Deco; Noisy Brain |
99 |
|
Inputs from the parietal cortex,
involved in spatial computation, project more to the dorsolateral prefrontal
cortex. |
|
2 |
Rolls
& Deco; Noisy Brain |
99 |
|
Inputs from the temporal lobe
visual cortex, involved in time dictations about objects, project
ventrolaterally to the prefrontal cortex. |
|
0 |
Rolls
& Deco; Noisy Brain |
99 |
|
It could be that the dorsolateral prefrontal cortex is
especially involved in spatial short-term memory, whereas the inferior convexity
prefrontal cortex more in object short-term memory. |
|
0 |
Rolls
& Deco; Noisy Brain |
99 |
|
The organization-by-stimulus-domain hypothesis holds that the spatial
('where') working memory is supported by the dorsolateral PFC, BA 46/9, in the middle frontal gyrus (MFG). |
|
0 |
Rolls
& Deco; Noisy Brain |
99 |
|
Object ('what') working memory is supported by the ventrolateral
PFC on the lateral/inferior
connectivity, BA 45. |
|
0 |
Rolls
& Deco; Noisy Brain |
99 |
|
Some mixing of the spatial and object representations in the lateral
prefrontal cortex is needed in order for some short-term memory tasks to be
resolved, and thus total segregation of 'what' and 'where' processing in the lateral prefrontal cortex should not be is expected. |
|
0 |
Rolls
& Deco; Noisy Brain |
|
|
|
|
|
|
|
|
|
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