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- A comparison of spike time prediction and receptive field mapping with point process generalized linear models, Wiener-Voltera kernels, and spike-triggered averaging methods (2009)
- Poster presentation: Characterizing neuronal encoding is essential for understanding information processing in the brain. Three methods are commonly used to characterize the relationship between neural spiking activity and the features of putative stimuli. These methods include: Wiener-Volterra kernel methods (WVK), the spike-triggered average (STA), and more recently, the point process generalized linear model (GLM). We compared the performance of these three approaches in estimating receptive field properties and orientation tuning of 251 V1 neurons recorded from 2 monkeys during a fixation period in response to a moving bar. The GLM consisted of two formulations of the conditional intensity function for a point process characterization of the spiking activity: one with a stimulus only component and one with the stimulus and spike history. We fit the GLMs by maximum likelihood using GLMfit in Matlab. Goodness-of-fit was assessed using cross-validation with Kolmogorov-Smirnov (KS) tests based on the time-rescaling theorem to evaluate the accuracy with which each model predicts the spiking activity of individual neurons and for each movement direction (4016 models in total, for 251 neurons and 16 different directions). The GLMs that considered spike history of up to 35 ms, accurately predicted neuronal spiking activity (95% confidence intervals for KS test) with a performance of 97.0% (3895/4016) for the training data, and 96.5% (3876/4016) for the test data. If spike history was not considered, performance dropped to 73,1% in the training and 71.3% in the testing data. In contrast, the WVF and the STA predicted spiking accurately for 24.2% and 44.5% of the test data examples respectively. The receptive field size estimates obtained from the GLM (with and without history), WVF and STA were comparable. Relative to the GLM orientation tuning was underestimated on average by a factor of 0.45 by the WVF and the STA. The main reason for using the STA and WVF approaches is their apparent simplicity. However, our analyses suggest that more accurate spike prediction as well as more credible estimates of receptive field size and orientation tuning can be computed easily using GLMs implemented in Matlab with standard functions such as GLMfit.

- Auto-structure of spike trains matters for testing on synchronous activity (2009)
- Poster presentation: Coordinated neuronal activity across many neurons, i.e. synchronous or spatiotemporal pattern, had been believed to be a major component of neuronal activity. However, the discussion if coordinated activity really exists remained heated and controversial. A major uncertainty was that many analysis approaches either ignored the auto-structure of the spiking activity, assumed a very simplified model (poissonian firing), or changed the auto-structure by spike jittering. We studied whether a statistical inference that tests whether coordinated activity is occurring beyond chance can be made false if one ignores or changes the real auto-structure of recorded data. To this end, we investigated the distribution of coincident spikes in mutually independent spike-trains modeled as renewal processes. We considered Gamma processes with different shape parameters as well as renewal processes in which the ISI distribution is log-normal. For Gamma processes of integer order, we calculated the mean number of coincident spikes, as well as the Fano factor of the coincidences, analytically. We determined how these measures depend on the bin width and also investigated how they depend on the firing rate, and on rate difference between the neurons. We used Monte-Carlo simulations to estimate the whole distribution for these parameters and also for other values of gamma. Moreover, we considered the effect of dithering for both of these processes and saw that while dithering does not change the average number of coincidences, it does change the shape of the coincidence distribution. Our major findings are: 1) the width of the coincidence count distribution depends very critically and in a non-trivial way on the detailed properties of the inter-spike interval distribution, 2) the dependencies of the Fano factor on the coefficient of variation of the ISI distribution are complex and mostly non-monotonic. Moreover, the Fano factor depends on the very detailed properties of the individual point processes, and cannot be predicted by the CV alone. Hence, given a recorded data set, the estimated value of CV of the ISI distribution is not sufficient to predict the Fano factor of the coincidence count distribution, and 3) spike jittering, even if it is as small as a fraction of the expected ISI, can falsify the inference on coordinated firing. In most of the tested cases and especially for complex synchronous and spatiotemporal pattern across many neurons, spike jittering increased the likelihood of false positive finding very strongly. Last, we discuss a procedure [1] that considers the complete auto-structure of each individual spike-train for testing whether synchrony firing occurs at chance and therefore overcomes the danger of an increased level of false positives.

- Detection of task-related synchronous firing patterns (2009)
- Poster presentation: Background To test the importance of synchronous neuronal firing for information processing in the brain, one has to investigate if synchronous firing strength is correlated to the experimental subjects. This requires a tool that can compare the strength of the synchronous firing across different conditions, while at the same time it should correct for other features of neuronal firing such as spike rate modulation or the auto-structure of the spike trains that might co-occur with synchronous firing. Here we present the bi- and multivariate extension of previously developed method NeuroXidence [1,2], which allows for comparing the amount of synchronous firing between different conditions. ...

- A mechanism for achieving zero-lag long-range synchronization of neural activity (2009)
- Poster presentation: How can two distant neural assemblies synchronize their firings at zero-lag even in the presence of non-negligible delays in the transfer of information between them? Neural synchronization stands today as one of the most promising mechanisms to counterbalance the huge anatomical and functional specialization of the different brain areas. However, and albeit more evidence is being accumulated in favor of its functional role as a binding mechanism of distributed neural responses, the physical and anatomical substrate for such a dynamic and precise synchrony, especially zero-lag even in the presence of non-negligible delays, remains unclear. Here we propose a simple network motif that naturally accounts for zero-lag synchronization of spiking assemblies of neurons for a wide range of temporal delays. We demonstrate that when two distant neural assemblies do not interact directly but relaying their dynamics via a third mediating single neuron or population and eventually achieve zero-lag coherent firing. Extensive numerical simulations of populations of Hodgkin-Huxley neurons interacting in such a network are analyzed. The results show that even with axonal delays as large as 15 ms the distant neural populations can synchronize their firings at zero-lag in a millisecond precision after the exchange of a few spikes. The role of noise and a distribution of axonal delays in the synchronized dynamics of the neural populations are also studied confirming the robustness of this sync mechanism. The proposed network module is densely embedded within the complex functional architecture of the brain and especially within the reciprocal thalamocortical interactions where the role of indirect pathways mimicking direct cortico-cortical fibers has been already suggested to facilitate trans-areal cortical communication. In summary the robust neural synchronization mechanism presented here arises as a consequence of the relay and redistribution of the dynamics performed by a mediating neuronal population. In opposition to previous works, neither inhibitory, gap junctions, nor complex networks need to be invoked to provide a stable mechanism of zero-phase correlated activity of neural populations in the presence of large conduction delays.

- Neural synchrony in cortical networks: history, concept and current status (2009)
- Following the discovery of context-dependent synchronization of oscillatory neuronal responses in the visual system, the role of neural synchrony in cortical networks has been expanded to provide a general mechanism for the coordination of distributed neural activity patterns. In the current paper, we present an update of the status of this hypothesis through summarizing recent results from our laboratory that suggest important new insights regarding the mechanisms, function and relevance of this phenomenon. In the first part, we present recent results derived from animal experiments and mathematical simulations that provide novel explanations and mechanisms for zero and nero-zero phase lag synchronization. In the second part, we shall discuss the role of neural synchrony for expectancy during perceptual organization and its role in conscious experience. This will be followed by evidence that indicates that in addition to supporting conscious cognition, neural synchrony is abnormal in major brain disorders, such as schizophrenia and autism spectrum disorders. We conclude this paper with suggestions for further research as well as with critical issues that need to be addressed in future studies.

- Context Matters: The Illusive Simplicity of Macaque V1 Receptive Fields (2012)
- Even in V1, where neurons have well characterized classical receptive fields (CRFs), it has been difficult to deduce which features of natural scenes stimuli they actually respond to. Forward models based upon CRF stimuli have had limited success in predicting the response of V1 neurons to natural scenes. As natural scenes exhibit complex spatial and temporal correlations, this could be due to surround effects that modulate the sensitivity of the CRF. Here, instead of attempting a forward model, we quantify the importance of the natural scenes surround for awake macaque monkeys by modeling it non-parametrically. We also quantify the influence of two forms of trial to trial variability. The first is related to the neuron’s own spike history. The second is related to ongoing mean field population activity reflected by the local field potential (LFP). We find that the surround produces strong temporal modulations in the firing rate that can be both suppressive and facilitative. Further, the LFP is found to induce a precise timing in spikes, which tend to be temporally localized on sharp LFP transients in the gamma frequency range. Using the pseudo R2 as a measure of model fit, we find that during natural scene viewing the CRF dominates, accounting for 60% of the fit, but that taken collectively the surround, spike history and LFP are almost as important, accounting for 40%. However, overall only a small proportion of V1 spiking statistics could be explained (R2~5%), even when the full stimulus, spike history and LFP were taken into account. This suggests that under natural scene conditions, the dominant influence on V1 neurons is not the stimulus, nor the mean field dynamics of the LFP, but the complex, incoherent dynamics of the network in which neurons are embedded.

- SORN: a self-organizing recurrent neural network (2009)
- Understanding the dynamics of recurrent neural networks is crucial for explaining how the brain processes information. In the neocortex, a range of different plasticity mechanisms are shaping recurrent networks into effective information processing circuits that learn appropriate representations for time-varying sensory stimuli. However, it has been difficult to mimic these abilities in artificial neural network models. Here we introduce SORN, a self-organizing recurrent network. It combines three distinct forms of local plasticity to learn spatio-temporal patterns in its input while maintaining its dynamics in a healthy regime suitable for learning. The SORN learns to encode information in the form of trajectories through its high-dimensional state space reminiscent of recent biological findings on cortical coding. All three forms of plasticity are shown to be essential for the network's success. Keywords: synaptic plasticity, intrinsic plasticity, recurrent neural networks, reservoir computing, time series prediction

- Transfer entropy - a model-free measure of effective connectivity for the neurosciences (2010)
- Understanding causal relationships, or effective connectivity, between parts of the brain is of utmost importance because a large part of the brain’s activity is thought to be internally generated and, hence, quantifying stimulus response relationships alone does not fully describe brain dynamics. Past efforts to determine effective connectivity mostly relied on model based approaches such as Granger causality or dynamic causal modeling. Transfer entropy (TE) is an alternative measure of effective connectivity based on information theory. TE does not require a model of the interaction and is inherently non-linear. We investigated the applicability of TE as a metric in a test for effective connectivity to electrophysiological data based on simulations and magnetoencephalography (MEG) recordings in a simple motor task. In particular, we demonstrate that TE improved the detectability of effective connectivity for non-linear interactions, and for sensor level MEG signals where linear methods are hampered by signal-cross-talk due to volume conduction.

- Spike train auto-structure impacts post-synaptic firing and timing-based plasticity (2011)
- Cortical neurons are typically driven by several thousand synapses. The precise spatiotemporal pattern formed by these inputs can modulate the response of a post-synaptic cell. In this work, we explore how the temporal structure of pre-synaptic inhibitory and excitatory inputs impact the post-synaptic firing of a conductance-based integrate and fire neuron. Both the excitatory and inhibitory input was modeled by renewal gamma processes with varying shape factors for modeling regular and temporally random Poisson activity. We demonstrate that the temporal structure of mutually independent inputs affects the post-synaptic firing, while the strength of the effect depends on the firing rates of both the excitatory and inhibitory inputs. In a second step, we explore the effect of temporal structure of mutually independent inputs on a simple version of Hebbian learning, i.e., hard bound spike-timing-dependent plasticity. We explore both the equilibrium weight distribution and the speed of the transient weight dynamics for different mutually independent gamma processes. We find that both the equilibrium distribution of the synaptic weights and the speed of synaptic changes are modulated by the temporal structure of the input. Finally, we highlight that the sensitivity of both the post-synaptic firing as well as the spike-timing-dependent plasticity on the auto-structure of the input of a neuron could be used to modulate the learning rate of synaptic modification.