Spike Correlations in a Songbird Agree with a Simple Markov Population Model

The relationships between neural activity at the single-cell and the population levels are of central importance for understanding neural codes. In many sensory systems, collective behaviors in large cell groups can be described by pairwise spike correlations. Here, we test whether in a highly specialized premotor system of songbirds, pairwise spike correlations themselves can be seen as a simple corollary of an underlying random process. We test hypotheses on connectivity and network dynamics in the motor pathway of zebra finches using a high-level population model that is independent of detailed single-neuron properties. We assume that neural population activity evolves along a finite set of states during singing, and that during sleep population activity randomly switches back and forth between song states and a single resting state. Individual spike trains are generated by associating with each of the population states a particular firing mode, such as bursting or tonic firing. With an overall modification of one or two simple control parameters, the Markov model is able to reproduce observed firing statistics and spike correlations in different neuron types and behavioral states. Our results suggest that song- and sleep-related firing patterns are identical on short time scales and result from random sampling of a unique underlying theme. The efficiency of our population model may apply also to other neural systems in which population hypotheses can be tested on recordings from small neuron groups

Unsupervised discovery of temporal sequences in high-dimensional datasets, with applications to neuroscience.

Identifying low-dimensional features that describe large-scale neural recordings is a major challenge in neuroscience. Repeated temporal patterns (sequences) are thought to be a salient feature of neural dynamics, but are not succinctly captured by traditional dimensionality reduction techniques. Here, we describe a software toolbox-called seqNMF-with new methods for extracting informative, non-redundant, sequences from high-dimensional neural data, testing the significance of these extracted patterns, and assessing the prevalence of sequential structure in data. We test these methods on simulated data under multiple noise conditions, and on several real neural and behavioral datas. In hippocampal data, seqNMF identifies neural sequences that match those calculated manually by reference to behavioral events. In songbird data, seqNMF discovers neural sequences in untutored birds that lack stereotyped songs. Thus, by identifying temporal structure directly from neural data, seqNMF enables dissection of complex neural circuits without relying on temporal references from stimuli or behavioral outputs

A reafferent and feed-forward model of song syntax generation in the Bengalese finch

Adult Bengalese finches generate a variable song that obeys a distinct and individual syntax. The syntax is gradually lost over a period of days after deafening and is recovered when hearing is restored. We present a spiking neuronal network model of the song syntax generation and its loss, based on the assumption that the syntax is stored in reafferent connections from the auditory to the motor control area. Propagating synfire activity in the HVC codes for individual syllables of the song and priming signals from the auditory network reduce the competition between syllables to allow only those transitions that are permitted by the syntax. Both imprinting of song syntax within HVC and the interaction of the reafferent signal with an efference copy of the motor command are sufficient to explain the gradual loss of syntax in the absence of auditory feedback. The model also reproduces for the first time experimental findings on the influence of altered auditory feedback on the song syntax generation, and predicts song- and species-specific low frequency components in the LFP. This study illustrates how sequential compositionality following a defined syntax can be realized in networks of spiking neurons

Methods for studying the neural code in high dimensions

Over the last two decades technological developments in multi-electrode arrays and fluorescence microscopy have made it possible to simultaneously record from hundreds to thousands of neurons. Developing methods for analyzing these data in order to learn how networks of neurons respond to external stimuli and process information is an outstanding challenge for neuroscience. In this dissertation, I address the challenge of developing and testing models that are both flexible and computationally tractable when used with high dimensional data. In chapter 2 I will discuss an approximation to the generalized linear model (GLM) log-likelihood that I developed in collaboration with my thesis advisor. This approximation is designed to ease the computational burden of evaluating GLMs. I will show that our method reduces the computational cost of evaluating the GLM log-likelihood by a factor proportional to the number of parameters in the model times the number of observations. Therefore it is most beneficial in typical neuroscience applications where the number of parameters is large. I then detail a variety of applications where our method can be of use, including Maximum Likelihood estimation of GLM parameters, marginal likelihood calculations for model selection and Markov chain Monte Carlo methods for sampling from posterior parameter distributions. I go on to show that our model does not necessarily sacrifice accuracy for speed. Using both analytic calculations and multi-unit, primate retinal responses, I show that parameter estimates and predictions using our model can have the same accuracy as that of generalized linear models. In chapter 3 I study the neural decoding problem of predicting stimuli from neuronal responses. The focus is on reconstructing zebra finch song spectrograms, which are high-dimensional, by combining the spike trains of zebra finch auditory midbrain neurons with information about the correlations present in all zebra finch song. I use a GLM to model neuronal responses and a series of prior distributions, each carrying different amounts of statistical information about zebra finch song. For song reconstruction I make use of recent connections made between the applied mathematics literature on solving linear systems of equations involving matrices with special structure and neural decoding. This allowed me to calculate \textit{maximum a posteriori} (MAP) estimates of song spectrograms in a time that only grows linearly, and is therefore quite tractable, with the number of time-bins in the song spectrogram. This speed was beneficial for answering questions which required the reconstruction of a variety of song spectrograms each corresponding to different priors made on the distribution of zebra finch song. My collaborators and I found that spike trains from a population of 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

Metastable attractors explain the variable timing of stable behavioral action sequences

info:eu-repo/semantics/publishedVersio

A multispecies perspective on ecological impacts of climatic forcing

P>1. In the prevailing context of concerns over climate change and its potential impacts on ecosystems, evaluating ecological consequences of climatic forcing has become a critical issue. 2. Historical data on the abundance of organisms have been extensively used to characterize the ecological effects of climatic forcing through specific weather and/or climatic variables, with most of the studies confined to single population models. 3. However, population responses to environmental fluctuations typically depend upon positive and negative feedbacks induced by interactions with other species. It is therefore important to integrate the insights gained from single population approaches into a multispecies perspective. 4. Here we combine the hierarchical Bayesian modelling approach with the state-space formulation to extend the scope of previously proposed models of population dynamics under climatic forcing to multi-species systems. 5. We use our model to analyse long-term macro-moth (Lepidoptera) community data from the Rothamsted Insect Survey network in the UK, using winter rainfall and winter temperature as environmental covariates. 6. The effects of the two weather variables were consistent across species, being negative for winter rainfall and positive for winter temperature. The two weather variables jointly explained 15-40% of the total environmental variation affecting the dynamics of individual species, and could explain up to 90% of covariances in species dynamics. 7. The contribution of interspecific interactions to community-level variation was found to be weak compared to the contributions of environmental forcing and intraspecific interactions

Bayesian multi-species modelling of non-negative continuous ecological data with a discrete mass at zero

Severe declines in the number of some songbirds over the last 40 years have caused heated debate amongst interested parties. Many factors have been suggested as possible causes for these declines, including an increase in the abundance and distribution of an avian predator, the Eurasian sparrowhawk Accipiter nisus. To test for evidence for a predator effect on the abundance of its prey, we analyse data on 10 species visiting garden bird feeding stations monitored by the British Trust for Ornithology in relation to the abundance of sparrowhawks. We apply Bayesian hierarchical models to data relating to averaged maximum weekly counts from a garden bird monitoring survey. These data are essentially continuous, bounded below by zero, but for many species show a marked spike at zero that many standard distributions would not be able to account for. We use the Tweedie distributions, which for certain areas of parameter space relate to continuous nonnegative distributions with a discrete probability mass at zero, and are hence able to deal with the shape of the empirical distributions of the data. The methods developed in this thesis begin by modelling single prey species independently with an avian predator as a covariate, using MCMC methods to explore parameter and model spaces. This model is then extended to a multiple-prey species model, testing for interactions between species as well as synchrony in their response to environmental factors and unobserved variation. Finally we use a relatively new methodological framework, namely the SPDE approach in the INLA framework, to fit a multi-species spatio-temporal model to the ecological data. The results from the analyses are consistent with the hypothesis that sparrowhawks are suppressing the numbers of some species of birds visiting garden feeding stations. Only the species most susceptible to sparrowhawk predation seem to be affected

Modeling and Analysis of Neural Spike Trains

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Hidden Markov Models

Hidden Markov Models (HMMs), although known for decades, have made a big career nowadays and are still in state of development. This book presents theoretical issues and a variety of HMMs applications in speech recognition and synthesis, medicine, neurosciences, computational biology, bioinformatics, seismology, environment protection and engineering. I hope that the reader will find this book useful and helpful for their own research