9 Organization Principles of Cortical Circuits and Their Function
isocortex hypothesis
Neocortex has relatively
uniform six-layered appeareance
Neocortex of all mammals is built of a common circuit
Composition
broadband transmission for local circuits in the gray matter
long-distance, low-bandwidth transmission for white matter
Pyramidal cells
80% of neruons
are spiny and have a prominent apical dendrite
Smooth cells
Spiny stellate
20% of neurons
do not have densely spiny dendrites
Glutamate neurotransmitter
excitatory
GABA neurotransmitter
inhibitory
spiny but lack of apical dendrite
exist only in middle layers of certain sensory cortical areas
axon of most cells does not project out of their own cortical area
axon of most cells does not project out of their own cortical area
Diversity
smooth cells have a variety of forms; pyramidal cells and spiny stellate cell differ in dendritic arborization and the precision of axonal projections
shape and size generates constraints on
- what connections are possible
- how many synapses they can support
Rules of engagement
Peters's Rule
neurons connect to each other in proportion to the amount of dendrite or axon they contribute to a particular layer
commonly interpreted as evidence for random connections
however, statistical averaging says nothing about specificity of neuron-to-neuron connections; quantification is needed
there are studies suggested specific connections exist while Peters's rule still holds in overall statistics
White's Exceptions
the deviation from Peters's Rule
Mapping on the body
distribution of excitatory and inhibitory neurons on their targets are nonrandom
recent studies shows that inhibitory synapses contact with all parts of the neuron
standard cable theory predicts that the most efficient site for inhibitor synapses is he proximal region around cell body, where indeed dominated by inhibitory neurons
inhibitory synapses at distal regions are found to be more efficient to quench distal excitatory signals
Synaptic connections
space-saving strategy for making most of connections with nearby neurons
long distance connections
long-distance connections in low in number
feedforward
feedback
number of neurons involved falls exponentially with distance
originate from neurons in superficial layers
contact to middle layer of the target area
contact outside the middle layers
originate from deep layers
lateral connections
originate from all layers
contact to all layers
Local processor
❗Recurrent circuits
a series of nested positive or negative feedback loops
Canonical circuit model
thalamic synapses are weak
there is strong recurrent excitation within and between layers
excitation and inhibition are linked in tandem to remain in balance and to prevent overexciting the circuit
express the functional relationships bw the excitatory and inhibitory neurons in the different cortical layers
show how the inputs to a local region of cortex from the sensory periphery via the thalamus or other cortical areas are integrated
explain how the relative tiny numbers of thalamic synapses are effective if they act in synchrony and are amplified by recurrent excitatory circuits
10 Cost, Efficiency, and Economy of Brain Networks
Cost
Efficiency
Wiring cost
Running cost
limits on brain size place constraints on the number and density of neurons
allometric scaling law
white matter volume growing faster than gray matter volume as brain size increases
spatially close neurons tend to be densely and mutually interconnected, conserving wiring cost
connection probabilities are reduced as physical distance increases
metabolic cost in maintaining brain function and its rest state
cause evolutional pressure to increases energy efficiency, neuronal coding strategy that minimize energy use with action potentials, and sparse coding
efficient network allows pairs of network elements to interact along relatively short or direct communication paths
small-world attributes
nodes are more likely to link with the neighboring nodes while having short path length to any nodes
network modules and hubs
each module corresponds to a set of densely interconnected nodes while connections bw modules are sparse
dense connections within modules and weak interconnections bw then for a structural basis for functional specialization
network hubs
specific subset of nodes dedicates to communication paths bw modules; act as points of convergence and divergence for intermodular signal
Economy
Pareto optimality
the construction or operation of a given system simultaneously satisfies multiple objectives
biological fitness or optimal design are derived from trade-off bw cost minimization and efficiency maximization
12 Mudularity, Plasticity, and Development of Brain Function
13 Experimential Effects in Neurocignitive Development
14 Plasticity as a Consequence of Experience and Aging
main idea
e.g. derease in GABAergic inhibition and increase spontaneous and driven neural activity
neuroplasticity is responsible for compensation mechanisms during natural aging
main idea
fitness is related to modularity of the brain, with highest fitness for low modularity in children, high modularity in young adult, decreased mudularity for elderly
main idea
plasiticity within a system exhibits both enhancements and deficits in processing
e.g. visual dorsal pathway exhibits high degree of plasticity compared to ventral path, visual tasks related to dorsal path can be enhanced (as in congenital deaf person) or impaired (as in dyslexic person)