Neuron as a reward-modulated combinatorial switch and a model of learning behavior

This paper proposes a neuronal circuitry layout and synaptic plasticity principles that allow the (pyramidal) neuron to act as a "combinatorial switch". Namely, the neuron learns to be more prone to generate spikes given those combinations of firing input neurons for which a previous spiking of the neuron had been followed by a positive global reward signal. The reward signal may be mediated by certain modulatory hormones or neurotransmitters, e.g., the dopamine. More generally, a trial-and-error learning paradigm is suggested in which a global reward signal triggers long-term enhancement or weakening of a neuron's spiking response to the preceding neuronal input firing pattern. Thus, rewards provide a feedback pathway that informs neurons whether their spiking was beneficial or detrimental for a particular input combination. The neuron's ability to discern specific combinations of firing input neurons is achieved through a random or predetermined spatial distribution of input synapses on dendrites that creates synaptic clusters that represent various permutations of input neurons. The corresponding dendritic segments, or the enclosed individual spines, are capable of being particularly excited, due to local sigmoidal thresholding involving voltage-gated channel conductances, if the segment's excitatory and absence of inhibitory inputs are temporally coincident. Such nonlinear excitation corresponds to a particular firing combination of input neurons, and it is posited that the excitation strength encodes the combinatorial memory and is regulated by long-term plasticity mechanisms. It is also suggested that the spine calcium influx that may result from the spatiotemporal synaptic input coincidence may cause the spine head actin filaments to undergo mechanical (muscle-like) contraction, with the ensuing cytoskeletal deformation transmitted to the axon initial segment where it may...Comment: Version 5: added computer code in the ancillary files sectio

An Operating Principle of the Cerebral Cortex, and a Cellular Mechanism for Attentional Trial-and-Error Pattern Learning and Useful Classification Extraction

A feature of the brains of intelligent animals is the ability to learn to respond to an ensemble of active neuronal inputs with a behaviorally appropriate ensemble of active neuronal outputs. Previously, a hypothesis was proposed on how this mechanism is implemented at the cellular level within the neocortical pyramidal neuron: the apical tuft or perisomatic inputs initiate "guess" neuron firings, while the basal dendrites identify input patterns based on excited synaptic clusters, with the cluster excitation strength adjusted based on reward feedback. This simple mechanism allows neurons to learn to classify their inputs in a surprisingly intelligent manner. Here, we revise and extend this hypothesis. We modify synaptic plasticity rules to align with behavioral time scale synaptic plasticity (BTSP) observed in hippocampal area CA1, making the framework more biophysically and behaviorally plausible. The neurons for the guess firings are selected in a voluntary manner via feedback connections to apical tufts in the neocortical layer 1, leading to dendritic Ca2+ spikes with burst firing, which are postulated to be neural correlates of attentional, aware processing. Once learned, the neuronal input classification is executed without voluntary or conscious control, enabling hierarchical incremental learning of classifications that is effective in our inherently classifiable world. In addition to voluntary, we propose that pyramidal neuron burst firing can be involuntary, also initiated via apical tuft inputs, drawing attention towards important cues such as novelty and noxious stimuli. We classify the excitations of neocortical pyramidal neurons into four categories based on their excitation pathway: attentional versus automatic and voluntary/acquired versus involuntary. Additionally, we hypothesize that dendrites within pyramidal neuron minicolumn bundles are coupled via depolarization...Comment: 20 pages, 13 figure

An operating principle of the cerebral cortex, and a cellular mechanism for attentional trial-and-error pattern learning and useful classification extraction

A feature of the brains of intelligent animals is the ability to learn to respond to an ensemble of active neuronal inputs with a behaviorally appropriate ensemble of active neuronal outputs. Previously, a hypothesis was proposed on how this mechanism is implemented at the cellular level within the neocortical pyramidal neuron: the apical tuft or perisomatic inputs initiate “guess” neuron firings, while the basal dendrites identify input patterns based on excited synaptic clusters, with the cluster excitation strength adjusted based on reward feedback. This simple mechanism allows neurons to learn to classify their inputs in a surprisingly intelligent manner. Here, we revise and extend this hypothesis. We modify synaptic plasticity rules to align with behavioral time scale synaptic plasticity (BTSP) observed in hippocampal area CA1, making the framework more biophysically and behaviorally plausible. The neurons for the guess firings are selected in a voluntary manner via feedback connections to apical tufts in the neocortical layer 1, leading to dendritic Ca2+ spikes with burst firing, which are postulated to be neural correlates of attentional, aware processing. Once learned, the neuronal input classification is executed without voluntary or conscious control, enabling hierarchical incremental learning of classifications that is effective in our inherently classifiable world. In addition to voluntary, we propose that pyramidal neuron burst firing can be involuntary, also initiated via apical tuft inputs, drawing attention toward important cues such as novelty and noxious stimuli. We classify the excitations of neocortical pyramidal neurons into four categories based on their excitation pathway: attentional versus automatic and voluntary/acquired versus involuntary. Additionally, we hypothesize that dendrites within pyramidal neuron minicolumn bundles are coupled via depolarization cross-induction, enabling minicolumn functions such as the creation of powerful hierarchical “hyperneurons” and the internal representation of the external world. We suggest building blocks to extend the microcircuit theory to network-level processing, which, interestingly, yields variants resembling the artificial neural networks currently in use. On a more speculative note, we conjecture that principles of intelligence in universes governed by certain types of physical laws might resemble ours

Relativistic description of 3He(e,e'p)2H

The Relativistic Distorted-Wave Impulse Approximation is used to describe the $^3$He($e,e^\prime p$)$^2$H process. We describe the $^3$He nucleus within the adiabatic hyperspherical expansion method with realistic nucleon-nucleon interactions. The overlap between the $^3$He and the deuteron wave functions can be accurately computed from a three-body calculation. The nucleons are described by solutions of the Dirac equation with scalar and vector (S-V) potentials. The wave function of the outgoing proton is obtained by solving the Dirac equation with a S-V optical potential fitted to elastic proton scattering data on the residual nucleus. Within this theoretical framework, we compute the cross section of the reaction and other observables like the transverse-longitudinal asymmetry, and compare them with the available experimental data measured at JLab.Comment: 4 pages, 3 figures. Proceedings of the 21st European Few Body Conference held in Salamanca (Spain) in August-September 201

The Quasielastic 3He(e,e'p)d Reaction at Q^2 = 1.5 GeV^2 for Recoil Momenta up to 1 GeV/c

We have studied the quasielastic 3He(e,e'p)d reaction in perpendicular coplanar kinematics, with the energy and momentum transferred by the electron fixed at 840 MeV and 1502 MeV/c, respectively. The 3He(e,e'p)d cross section was measured for missing momenta up to 1000 MeV/c, while the A_TL asymmetry was extracted for missing momenta up to 660 MeV/c. For missing momenta up to 150 MeV/c, the measured cross section is described well by calculations that use a variational ground-state wave function of the 3He nucleus derived from a potential that includes three-body forces. For missing momenta from 150 to 750 MeV/c, strong final-state interaction effects are observed. Near 1000 MeV/c, the experimental cross section is more than an order of magnitude larger than predicted by available theories. The A_TL asymmetry displays characteristic features of broken factorization, and is described reasonably well by available models.Comment: 5 pages, 3 figures, submitted to Physical Review Letters, v3: changed conten

Recoil Polarization Measurements for Neutral Pion Electroproduction at Q^2=1 (GeV/c)^2 Near the Delta Resonance

We measured angular distributions of differential cross section, beam analyzing power, and recoil polarization for neutral pion electroproduction at Q^2 = 1.0 (GeV/c)^2 in 10 bins of W across the Delta resonance. A total of 16 independent response functions were extracted, of which 12 were observed for the first time. Comparisons with recent model calculations show that response functions governed by real parts of interference products are determined relatively well near 1.232 GeV, but variations among models is large for response functions governed by imaginary parts and for both increases rapidly with W. We performed a nearly model-independent multipole analysis that adjusts complex multipoles with high partial waves constrained by baseline models. Parabolic fits to the W dependence of the multipole analysis around the Delta mass gives values for SMR = (-6.61 +/- 0.18)% and EMR = (-2.87 +/- 0.19)% that are distinctly larger than those from Legendre analysis of the same data. Similarly, the multipole analysis gives Re(S0+/M1+) = (+7.1 +/- 0.8)% at W=1.232 GeV, consistent with recent models, while the traditional Legendre analysis gives the opposite sign because its truncation errors are quite severe. Finally, using a unitary isobar model (UIM), we find that excitation of the Roper resonance is dominantly longitudinal with S1/2 = (0.05 +/- 0.01) GeV^(-1/2) at Q^2=1. The ReS0+ and ReE0+ multipoles favor pseudovector coupling over pseudoscalar coupling or a recently proposed mixed-coupling scheme, but the UIM does not reproduce the imaginary parts of 0+ multipoles well.Comment: 60 pages, 54 figure

Phase transitions in contagion processes mediated by recurrent mobility patterns

Human mobility and activity patterns mediate contagion on many levels, including the spatial spread of infectious diseases, diffusion of rumors, and emergence of consensus. These patterns however are often dominated by specific locations and recurrent flows and poorly modeled by the random diffusive dynamics generally used to study them. Here we develop a theoretical framework to analyze contagion within a network of locations where individuals recall their geographic origins. We find a phase transition between a regime in which the contagion affects a large fraction of the system and one in which only a small fraction is affected. This transition cannot be uncovered by continuous deterministic models due to the stochastic features of the contagion process and defines an invasion threshold that depends on mobility parameters, providing guidance for controlling contagion spread by constraining mobility processes. We recover the threshold behavior by analyzing diffusion processes mediated by real human commuting data.Comment: 20 pages of Main Text including 4 figures, 7 pages of Supplementary Information; Nature Physics (2011

Measurement of the 3He(e,e'p)pn reaction at high missing energies and momenta

Results of the Jefferson Lab Hall A quasielastic 3He(e,e'p)pn measurements are presented. These measurements were performed at fixed transferred momentum and energy, q = 1502 MeV/c and omega = 840 MeV, respectively, for missing momenta p_m up to 1 GeV/c and missing energies in the continuum region, up to pion threshold; this kinematic coverage is much more extensive than that of any previous experiment. The cross section data are presented along with the effective momentum density distribution and compared to theoretical models.Comment: 5 pages, 3 figures, updated to reflect published paper: minor text changes from previous version along with updated and added reference