Synaptic shot noise and conductance fluctuations affect the membrane voltage with equal significance

The subthresholdmembranevoltage of a neuron in active cortical tissue is a fluctuating quantity with a distribution that reflects the firing statistics of the presynaptic population. It was recently found that conductancebased synaptic drive can lead to distributions with a significant skew. Here it is demonstrated that the underlying shot noise caused by Poissonian spike arrival also skews the membrane distribution, but in the opposite sense. Using a perturbative method, we analyze the effects of shot noise on the distribution of synaptic conductances and calculate the consequent voltage distribution. To first order in the perturbation theory, the voltage distribution is a gaussian modulated by a prefactor that captures the skew. The gaussian component is identical to distributions derived using current-based models with an effective membrane time constant. The well-known effective-time-constant approximation can therefore be identified as the leading-order solution to the full conductance-based model. The higher-order modulatory prefactor containing the skew comprises terms due to both shot noise and conductance fluctuations. The diffusion approximation misses these shot-noise effects implying that analytical approaches such as the Fokker-Planck equation or simulation with filtered white noise cannot be used to improve on the gaussian approximation. It is further demonstrated that quantities used for fitting theory to experiment, such as the voltage mean and variance, are robust against these non-Gaussian effects. The effective-time-constant approximation is therefore relevant to experiment and provides a simple analytic base on which other pertinent biological details may be added

Signal buffering in random networks of spiking neurons: microscopic vs. macroscopic phenomena

In randomly connected networks of pulse-coupled elements a time-dependent input signal can be buffered over a short time. We studied the signal buffering properties in simulated networks as a function of the networks state, characterized by both the Lyapunov exponent of the microscopic dynamics and the macroscopic activity derived from mean-field theory. If all network elements receive the same signal, signal buffering over delays comparable to the intrinsic time constant of the network elements can be explained by macroscopic properties and works best at the phase transition to chaos. However, if only 20 percent of the network units receive a common time-dependent signal, signal buffering properties improve and can no longer be attributed to the macroscopic dynamics.Comment: 5 pages, 3 figure

Addressing serial-order and negative-keying effects: A mixed-methods study

Researchers have studied item serial-order effects on attitudinal instruments by considering how item-total correlations differ based on the item’s placement within a scale (e.g., Hamilton & Shuminsky, 1990). In addition, other researchers have focused on item negative-keying effects on attitudinal instruments (e.g., Marsh, 1996). Researchers consistently have found that negatively-keyed items relate to one another above and beyond their relationship to the construct intended to be measured. However, only one study (i.e., Bandalos & Coleman, 2012) investigated the combined effects of serial-order and negative-keying on attitudinal instruments. Their brief study found some improvements in fit when attitudinal items were presented in a unique, random order to each participant, which is easily implemented using computer survey software. In this study I replicated and extended these findings by considering three attitudinal scales: Conformity Scale (Goldberg et al., 2006; Jackson, 1994) and two subscales of the Big Five – Conscientiousness and Agreeableness (John & Srivastava, 1999). In addition, I collected and analyzed qualitative data in the form of think-alouds and used these data to inform the quantitative results in an explanatory sequential mixed-methods design (Creswell, 2011). I administered three different groupings of the items on these three instruments to random groups of university students. The items were displayed in either a blocked (i.e., all positively-keyed items followed by all negatively-keyed items), alternating (i.e., items alternated keying every other item beginning with a positively-keyed item), or random (i.e., items presented in a different random order for each participant) order. When each participant saw a different randomly-ordered version of the attitudinal scale, I found fewer expected measurement error correlations among items of the same keying and in close proximity (i.e., serial order) to one another. Moreover, in this random ordering, the modification indices associated with the suggested measurement error correlations were lower than in the other orderings. Finally, the fit of the model to the data was the best in the random ordering for all except the Agreeableness scale. Practitioners are urged to administer attitudinal scales in a computer-generated random order unique to each participant whenever possible

Extracting non-linear integrate-and-fire models from experimental data using dynamic I–V curves

The dynamic I–V curve method was recently introduced for the efficient experimental generation of reduced neuron models. The method extracts the response properties of a neuron while it is subject to a naturalistic stimulus that mimics in vivo-like fluctuating synaptic drive. The resulting history-dependent, transmembrane current is then projected onto a one-dimensional current–voltage relation that provides the basis for a tractable non-linear integrate-and-fire model. An attractive feature of the method is that it can be used in spike-triggered mode to quantify the distinct patterns of post-spike refractoriness seen in different classes of cortical neuron. The method is first illustrated using a conductance-based model and is then applied experimentally to generate reduced models of cortical layer-5 pyramidal cells and interneurons, in injected-current and injected- conductance protocols. The resulting low-dimensional neuron models—of the refractory exponential integrate-and-fire type—provide highly accurate predictions for spike-times. The method therefore provides a useful tool for the construction of tractable models and rapid experimental classification of cortical neurons

Quaternion Singular Value Decomposition based on Bidiagonalization to a Real Matrix using Quaternion Householder Transformations

We present a practical and efficient means to compute the singular value decomposition (svd) of a quaternion matrix A based on bidiagonalization of A to a real bidiagonal matrix B using quaternionic Householder transformations. Computation of the svd of B using an existing subroutine library such as lapack provides the singular values of A. The singular vectors of A are obtained trivially from the product of the Householder transformations and the real singular vectors of B. We show in the paper that left and right quaternionic Householder transformations are different because of the noncommutative multiplication of quaternions and we present formulae for computing the Householder vector and matrix in each case

Noise Induced Coherence in Neural Networks

We investigate numerically the dynamics of large networks of $N$ globally pulse-coupled integrate and fire neurons in a noise-induced synchronized state. The powerspectrum of an individual element within the network is shown to exhibit in the thermodynamic limit ($N\to \infty$) a broadband peak and an additional delta-function peak that is absent from the powerspectrum of an isolated element. The powerspectrum of the mean output signal only exhibits the delta-function peak. These results are explained analytically in an exactly soluble oscillator model with global phase coupling.Comment: 4 pages ReVTeX and 3 postscript figure

Stability of Negative Image Equilibria in Spike-Timing Dependent Plasticity

We investigate the stability of negative image equilibria in mean synaptic weight dynamics governed by spike-timing dependent plasticity (STDP). The neural architecture of the model is based on the electrosensory lateral line lobe (ELL) of mormyrid electric fish, which forms a negative image of the reafferent signal from the fish's own electric discharge to optimize detection of external electric fields. We derive a necessary and sufficient condition for stability, for arbitrary postsynaptic potential functions and arbitrary learning rules. We then apply the general result to several examples of biological interest.Comment: 13 pages, revtex4; uses packages: graphicx, subfigure; 9 figures, 16 subfigure

Supervised Learning in Multilayer Spiking Neural Networks

The current article introduces a supervised learning algorithm for multilayer spiking neural networks. The algorithm presented here overcomes some limitations of existing learning algorithms as it can be applied to neurons firing multiple spikes and it can in principle be applied to any linearisable neuron model. The algorithm is applied successfully to various benchmarks, such as the XOR problem and the Iris data set, as well as complex classifications problems. The simulations also show the flexibility of this supervised learning algorithm which permits different encodings of the spike timing patterns, including precise spike trains encoding.Comment: 38 pages, 4 figure

Event-driven simulations of a plastic, spiking neural network

We consider a fully-connected network of leaky integrate-and-fire neurons with spike-timing-dependent plasticity. The plasticity is controlled by a parameter representing the expected weight of a synapse between neurons that are firing randomly with the same mean frequency. For low values of the plasticity parameter, the activities of the system are dominated by noise, while large values of the plasticity parameter lead to self-sustaining activity in the network. We perform event-driven simulations on finite-size networks with up to 128 neurons to find the stationary synaptic weight conformations for different values of the plasticity parameter. In both the low and high activity regimes, the synaptic weights are narrowly distributed around the plasticity parameter value consistent with the predictions of mean-field theory. However, the distribution broadens in the transition region between the two regimes, representing emergent network structures. Using a pseudophysical approach for visualization, we show that the emergent structures are of "path" or "hub" type, observed at different values of the plasticity parameter in the transition region.Comment: 9 pages, 6 figure

Adaptation Reduces Variability of the Neuronal Population Code

Sequences of events in noise-driven excitable systems with slow variables often show serial correlations among their intervals of events. Here, we employ a master equation for general non-renewal processes to calculate the interval and count statistics of superimposed processes governed by a slow adaptation variable. For an ensemble of spike-frequency adapting neurons this results in the regularization of the population activity and an enhanced post-synaptic signal decoding. We confirm our theoretical results in a population of cortical neurons.Comment: 4 pages, 2 figure