Assistant Professor, Department of Biology
Ph.D. Columbia University
B.Sc. Sharif University of Technology, Tehran, Iran
Research Interests: Theoretical neuroscience
Overview: Our lab's research is in theoretical neuroscience. Our broad interest is in understanding how large networks of neurons, e.g. those in the mammalian cerebral cortex, process sensory inputs and give rise to higher-level cognitive functions through their collective dynamics on multiple time scales. To shed light on the complexity of neurobiological phenomena we use mathematical models that capture a few core concepts or computational and dynamical principles. We also work on developing new statistical and computational tools for analyzing large, high-dimensional neurobiological and behavioral datasets. In pursuing these goals we use techniques from statistical physics, random matrix theory, machine learning and information theory. We collaborate with experimental labs here in the University of Oregon and elsewhere.
Current questions of interest include the following. How do randomness and nonnormality in the connectivity structure of networks affect their dynamics? What roles do the horizontal and feedback connections in sensory cortical areas play in contextual modulation (how e.g. the response of neurons in the visual cortex is affected by the visual context surrounding that stimulus) and ultimately in the dynamical representation of objects? Can the breakup of neural response types in the early auditory system be explained by efficient coding principles?
Learning unbelievable probabilities.
Adv Neural Inf Process Syst. 2011 Dec;24:738-746
Authors: Pitkow X, Ahmadian Y, Miller KD
Loopy belief propagation performs approximate inference on graphical models with loops. One might hope to compensate for the approximation by adjusting model parameters. Learning algorithms for this purpose have been explored previously, and the claim has been made that every set of locally consistent marginals can arise from belief propagation run on a graphical model. On the contrary, here we show that many probability distributions have marginals that cannot be reached by belief propagation using any set of model parameters or any learning algorithm. We call such marginals 'unbelievable.' This problem occurs whenever the Hessian of the Bethe free energy is not positive-definite at the target marginals. All learning algorithms for belief propagation necessarily fail in these cases, producing beliefs or sets of beliefs that may even be worse than the pre-learning approximation. We then show that averaging inaccurate beliefs, each obtained from belief propagation using model parameters perturbed about some learned mean values, can achieve the unbelievable marginals.
PMID: 28781497 [PubMed]
Properties of networks with partially structured and partially random connectivity.
Phys Rev E Stat Nonlin Soft Matter Phys. 2015 Jan;91(1):012820
Authors: Ahmadian Y, Fumarola F, Miller KD
Networks studied in many disciplines, including neuroscience and mathematical biology, have connectivity that may be stochastic about some underlying mean connectivity represented by a non-normal matrix. Furthermore, the stochasticity may not be independent and identically distributed (iid) across elements of the connectivity matrix. More generally, the problem of understanding the behavior of stochastic matrices with nontrivial mean structure and correlations arises in many settings. We address this by characterizing large random N×N matrices of the form A=M+LJR, where M,L, and R are arbitrary deterministic matrices and J is a random matrix of zero-mean iid elements. M can be non-normal, and L and R allow correlations that have separable dependence on row and column indices. We first provide a general formula for the eigenvalue density of A. For A non-normal, the eigenvalues do not suffice to specify the dynamics induced by A, so we also provide general formulas for the transient evolution of the magnitude of activity and frequency power spectrum in an N-dimensional linear dynamical system with a coupling matrix given by A. These quantities can also be thought of as characterizing the stability and the magnitude of the linear response of a nonlinear network to small perturbations about a fixed point. We derive these formulas and work them out analytically for some examples of M,L, and R motivated by neurobiological models. We also argue that the persistence as N→∞ of a finite number of randomly distributed outlying eigenvalues outside the support of the eigenvalue density of A, as previously observed, arises in regions of the complex plane Ω where there are nonzero singular values of L(-1)(z1-M)R(-1) (for z∈Ω) that vanish as N→∞. When such singular values do not exist and L and R are equal to the identity, there is a correspondence in the normalized Frobenius norm (but not in the operator norm) between the support of the spectrum of A for J of norm σ and the σ pseudospectrum of M.
PMID: 25679669 [PubMed - indexed for MEDLINE]
Analysis of the stabilized supralinear network.
Neural Comput. 2013 Aug;25(8):1994-2037
Authors: Ahmadian Y, Rubin DB, Miller KD
We study a rate-model neural network composed of excitatory and inhibitory neurons in which neuronal input-output functions are power laws with a power greater than 1, as observed in primary visual cortex. This supralinear input-output function leads to supralinear summation of network responses to multiple inputs for weak inputs. We show that for stronger inputs, which would drive the excitatory subnetwork to instability, the network will dynamically stabilize provided feedback inhibition is sufficiently strong. For a wide range of network and stimulus parameters, this dynamic stabilization yields a transition from supralinear to sublinear summation of network responses to multiple inputs. We compare this to the dynamic stabilization in the balanced network, which yields only linear behavior. We more exhaustively analyze the two-dimensional case of one excitatory and one inhibitory population. We show that in this case, dynamic stabilization will occur whenever the determinant of the weight matrix is positive and the inhibitory time constant is sufficiently small, and analyze the conditions for supersaturation, or decrease of firing rates with increasing stimulus contrast (which represents increasing input firing rates). In work to be presented elsewhere, we have found that this transition from supralinear to sublinear summation can explain a wide variety of nonlinearities in cerebral cortical processing.
PMID: 23663149 [PubMed - indexed for MEDLINE]
Modeling the impact of common noise inputs on the network activity of retinal ganglion cells.
J Comput Neurosci. 2012 Aug;33(1):97-121
Authors: Vidne M, Ahmadian Y, Shlens J, Pillow JW, Kulkarni J, Litke AM, Chichilnisky EJ, Simoncelli E, Paninski L
Synchronized spontaneous firing among retinal ganglion cells (RGCs), on timescales faster than visual responses, has been reported in many studies. Two candidate mechanisms of synchronized firing include direct coupling and shared noisy inputs. In neighboring parasol cells of primate retina, which exhibit rapid synchronized firing that has been studied extensively, recent experimental work indicates that direct electrical or synaptic coupling is weak, but shared synaptic input in the absence of modulated stimuli is strong. However, previous modeling efforts have not accounted for this aspect of firing in the parasol cell population. Here we develop a new model that incorporates the effects of common noise, and apply it to analyze the light responses and synchronized firing of a large, densely-sampled network of over 250 simultaneously recorded parasol cells. We use a generalized linear model in which the spike rate in each cell is determined by the linear combination of the spatio-temporally filtered visual input, the temporally filtered prior spikes of that cell, and unobserved sources representing common noise. The model accurately captures the statistical structure of the spike trains and the encoding of the visual stimulus, without the direct coupling assumption present in previous modeling work. Finally, we examined the problem of decoding the visual stimulus from the spike train given the estimated parameters. The common-noise model produces Bayesian decoding performance as accurate as that of a model with direct coupling, but with significantly more robustness to spike timing perturbations.
PMID: 22203465 [PubMed - indexed for MEDLINE]
Designing optimal stimuli to control neuronal spike timing.
J Neurophysiol. 2011 Aug;106(2):1038-53
Authors: Ahmadian Y, Packer AM, Yuste R, Paninski L
Recent advances in experimental stimulation methods have raised the following important computational question: how can we choose a stimulus that will drive a neuron to output a target spike train with optimal precision, given physiological constraints? Here we adopt an approach based on models that describe how a stimulating agent (such as an injected electrical current or a laser light interacting with caged neurotransmitters or photosensitive ion channels) affects the spiking activity of neurons. Based on these models, we solve the reverse problem of finding the best time-dependent modulation of the input, subject to hardware limitations as well as physiologically inspired safety measures, that causes the neuron to emit a spike train that with highest probability will be close to a target spike train. We adopt fast convex constrained optimization methods to solve this problem. Our methods can potentially be implemented in real time and may also be generalized to the case of many cells, suitable for neural prosthesis applications. With the use of biologically sensible parameters and constraints, our method finds stimulation patterns that generate very precise spike trains in simulated experiments. We also tested the intracellular current injection method on pyramidal cells in mouse cortical slices, quantifying the dependence of spiking reliability and timing precision on constraints imposed on the applied currents.
PMID: 21511704 [PubMed - indexed for MEDLINE]
Incorporating naturalistic correlation structure improves spectrogram reconstruction from neuronal activity in the songbird auditory midbrain.
J Neurosci. 2011 Mar 9;31(10):3828-42
Authors: Ramirez AD, Ahmadian Y, Schumacher J, Schneider D, Woolley SM, Paninski L
Birdsong is comprised of rich spectral and temporal organization, which might be used for vocal perception. To quantify how this structure could be used, we have reconstructed birdsong spectrograms by combining the spike trains of zebra finch auditory midbrain neurons with information about the correlations present in song. We calculated maximum a posteriori estimates of song spectrograms using a generalized linear model of neuronal responses and a series of prior distributions, each carrying different amounts of statistical information about zebra finch song. We found that spike trains from a population of mesencephalicus lateral dorsalis (MLd) neurons combined with an uncorrelated Gaussian prior can estimate the amplitude envelope of song spectrograms. The same set of responses can be combined with Gaussian priors that have correlations matched to those found across multiple zebra finch songs to yield song spectrograms similar to those presented to the animal. The fidelity of spectrogram reconstructions from MLd responses relies more heavily on prior knowledge of spectral correlations than temporal correlations. However, the best reconstructions combine MLd responses with both spectral and temporal correlations.
PMID: 21389238 [PubMed - indexed for MEDLINE]
Efficient Markov chain Monte Carlo methods for decoding neural spike trains.
Neural Comput. 2011 Jan;23(1):46-96
Authors: Ahmadian Y, Pillow JW, Paninski L
Stimulus reconstruction or decoding methods provide an important tool for understanding how sensory and motor information is represented in neural activity. We discuss Bayesian decoding methods based on an encoding generalized linear model (GLM) that accurately describes how stimuli are transformed into the spike trains of a group of neurons. The form of the GLM likelihood ensures that the posterior distribution over the stimuli that caused an observed set of spike trains is log concave so long as the prior is. This allows the maximum a posteriori (MAP) stimulus estimate to be obtained using efficient optimization algorithms. Unfortunately, the MAP estimate can have a relatively large average error when the posterior is highly nongaussian. Here we compare several Markov chain Monte Carlo (MCMC) algorithms that allow for the calculation of general Bayesian estimators involving posterior expectations (conditional on model parameters). An efficient version of the hybrid Monte Carlo (HMC) algorithm was significantly superior to other MCMC methods for gaussian priors. When the prior distribution has sharp edges and corners, on the other hand, the "hit-and-run" algorithm performed better than other MCMC methods. Using these algorithms, we show that for this latter class of priors, the posterior mean estimate can have a considerably lower average error than MAP, whereas for gaussian priors, the two estimators have roughly equal efficiency. We also address the application of MCMC methods for extracting nonmarginal properties of the posterior distribution. For example, by using MCMC to calculate the mutual information between the stimulus and response, we verify the validity of a computationally efficient Laplace approximation to this quantity for gaussian priors in a wide range of model parameters; this makes direct model-based computation of the mutual information tractable even in the case of large observed neural populations, where methods based on binning the spike train fail. Finally, we consider the effect of uncertainty in the GLM parameters on the posterior estimators.
PMID: 20964539 [PubMed - indexed for MEDLINE]
Model-based decoding, information estimation, and change-point detection techniques for multineuron spike trains.
Neural Comput. 2011 Jan;23(1):1-45
Authors: Pillow JW, Ahmadian Y, Paninski L
One of the central problems in systems neuroscience is to understand how neural spike trains convey sensory information. Decoding methods, which provide an explicit means for reading out the information contained in neural spike responses, offer a powerful set of tools for studying the neural coding problem. Here we develop several decoding methods based on point-process neural encoding models, or forward models that predict spike responses to stimuli. These models have concave log-likelihood functions, which allow efficient maximum-likelihood model fitting and stimulus decoding. We present several applications of the encoding model framework to the problem of decoding stimulus information from population spike responses: (1) a tractable algorithm for computing the maximum a posteriori (MAP) estimate of the stimulus, the most probable stimulus to have generated an observed single- or multiple-neuron spike train response, given some prior distribution over the stimulus; (2) a gaussian approximation to the posterior stimulus distribution that can be used to quantify the fidelity with which various stimulus features are encoded; (3) an efficient method for estimating the mutual information between the stimulus and the spike trains emitted by a neural population; and (4) a framework for the detection of change-point times (the time at which the stimulus undergoes a change in mean or variance) by marginalizing over the posterior stimulus distribution. We provide several examples illustrating the performance of these estimators with simulated and real neural data.
PMID: 20964538 [PubMed - indexed for MEDLINE]