Introducing a New Mobile Electroencephalography System and Evaluating Its Quality in Comparison to Clinical Electroencephalography.

Electroencephalography (EEG) is a crucial tool in cognitive neuroscience, enabling the study of neurophysiological function by measuring the brain's electrical activity. Its applications include perception, learning, memory, language, decision making and neural network mapping. Recently, interest has surged in extending EEG measurements to domestic environments. However, the high costs associated with traditional laboratory EEG systems have hindered accessibility for many individuals and researchers in education, research, and medicine. To tackle this, a mobile-EEG device named "DreamMachine" was developed. A more affordable alternative to both lab-based EEG systems and existing mobile-EEG devices. This system boasts 24 channels, 24-bit resolution, up to 6 h of battery life, portability, and a low price. Our open-source and open-hardware approach empowers cognitive neuroscience, especially in education, learning, and research, opening doors to more accessibility. This paper introduces the DreamMachine's design and compares it with the lab-based EEG system "asalabTM" in an eyes-open and eyes-closed experiment. The Alpha band exhibited higher power in the power spectrum during eyes-closed conditions, whereas the eyes-open condition showed increased power specifically within the Delta frequency range. Our analysis confirms that the DreamMachine accurately records brain activity, meeting the necessary standards when compared to the asalabTM system.

Learning in the Single-Cell Organism Physarum polycephalum: Effect of Propofol.

Propofol belongs to a class of molecules that are known to block learning and memory in mammals, including rodents and humans. Interestingly, learning and memory are not tied to the presence of a nervous system. There are several lines of evidence indicating that single-celled organisms also have the capacity for learning and memory which may be considered as basal intelligence. Here, we introduce a new experimental model for testing the learning ability of Physarum polycephalum, a model organism frequently used to study single-celled "intelligence". In this study, the impact of propofol on Physarum's "intelligence" was tested. The model consists of a labyrinth of subsequent bifurcations in which food (oat flakes soaked with coconut oil-derived medium chain triglycerides [MCT] and soybean oil-derived long chain triglycerides [LCT]) or propofol in MCT/LCT) is placed in one of each Y-branch. In this setting, it was tested whether Physarum memorized the rewarding branch. We saw that Physarum was a quick learner when capturing the first bifurcations of the maze; thereafter, the effect decreased, perhaps due to reaching a state of satiety. In contrast, when oat flakes were soaked with propofol, Physarum's preference for oat flakes declined significantly. Several possible actions, including the blocking of gamma-aminobutyric acid (GABA) receptor signaling, are suggested to account for this behavior, many of which can be tested in our new model.

Westdrive X LoopAR: An Open-Access Virtual Reality Project in Unity for Evaluating User Interaction Methods during Takeover Requests.

With the further development of highly automated vehicles, drivers will engage in non-related tasks while being driven. Still, drivers have to take over control when requested by the car. Here, the question arises, how potentially distracted drivers get back into the control-loop quickly and safely when the car requests a takeover. To investigate effective human-machine interactions, a mobile, versatile, and cost-efficient setup is needed. Here, we describe a virtual reality toolkit for the Unity 3D game engine containing all the necessary code and assets to enable fast adaptations to various human-machine interaction experiments, including closely monitoring the subject. The presented project contains all the needed functionalities for realistic traffic behavior, cars, pedestrians, and a large, open-source, scriptable, and modular VR environment. It covers roughly 25 km2, a package of 125 animated pedestrians, and numerous vehicles, including motorbikes, trucks, and cars. It also contains all the needed nature assets to make it both highly dynamic and realistic. The presented repository contains a C++ library made for LoopAR that enables force feedback for gaming steering wheels as a fully supported component. It also includes all necessary scripts for eye-tracking in the used devices. All the main functions are integrated into the graphical user interface of the Unity® editor or are available as prefab variants to ease the use of the embedded functionalities. This project's primary purpose is to serve as an open-access, cost-efficient toolkit that enables interested researchers to conduct realistic virtual reality research studies without costly and immobile simulators. To ensure the accessibility and usability of the mentioned toolkit, we performed a user experience report, also included in this paper.

Reliability and comparability of human brain structural covariance networks.

Structural covariance analysis is a widely used structural MRI analysis method which characterises the co-relations of morphology between brain regions over a group of subjects. To our knowledge, little has been investigated in terms of the comparability of results between different data sets of healthy human subjects, as well as the reliability of results over the same subjects in different rescan sessions, image resolutions, or FreeSurfer versions. In terms of comparability, our results show substantial differences in the structural covariance matrix between data sets of age- and sex-matched healthy human adults. These differences persist after univariate site correction, they are exacerbated by low sample sizes, and they are most pronounced when using average cortical thickness as a morphological measure. Down-stream graph theoretic analyses further show statistically significant differences. In terms of reliability, substantial differences were also found when comparing repeated scan sessions of the same subjects, image resolutions, and even FreeSurfer versions of the same image. We could further estimate the relative measurement error and showed that it is largest when using cortical thickness as a morphological measure. Using simulated data, we argue that cortical thickness is least reliable because of larger relative measurement errors. Practically, we make the following recommendations (1) combining subjects across sites into one group should be avoided, particularly if sites differ in image resolutions, subject demographics, or preprocessing steps; (2) surface area and volume should be preferred as morphological measures over cortical thickness; (3) a large number of subjects (n≫30 for the Desikan-Killiany parcellation) should be used to estimate structural covariance; (4) measurement error should be assessed where repeated measurements are available; (5) if combining sites is critical, univariate (per ROI) site-correction is insufficient, but error covariance (between ROIs) should be explicitly measured and modelled.

Human Decisions in Moral Dilemmas are Largely Described by Utilitarianism: Virtual Car Driving Study Provides Guidelines for Autonomous Driving Vehicles.

Ethical thought experiments such as the trolley dilemma have been investigated extensively in the past, showing that humans act in utilitarian ways, trying to cause as little overall damage as possible. These trolley dilemmas have gained renewed attention over the past few years, especially due to the necessity of implementing moral decisions in autonomous driving vehicles (ADVs). We conducted a set of experiments in which participants experienced modified trolley dilemmas as drivers in virtual reality environments. Participants had to make decisions between driving in one of two lanes where different obstacles came into view. Eventually, the participants had to decide which of the objects they would crash into. Obstacles included a variety of human-like avatars of different ages and group sizes. Furthermore, the influence of sidewalks as potential safe harbors and a condition implicating self-sacrifice were tested. Results showed that participants, in general, decided in a utilitarian manner, sparing the highest number of avatars possible with a limited influence by the other variables. Derived from these findings, which are in line with the utilitarian approach in moral decision making, it will be argued for an obligatory ethics setting implemented in ADVs.

A Unifying Framework of Synaptic and Intrinsic Plasticity in Neural Populations.

A neuronal population is a computational unit that receives a multivariate, time-varying input signal and creates a related multivariate output. These neural signals are modeled as stochastic processes that transmit information in real time, subject to stochastic noise. In a stationary environment, where the input signals can be characterized by constant statistical properties, the systematic relationship between its input and output processes determines the computation carried out by a population. When these statistical characteristics unexpectedly change, the population needs to adapt to its new environment if it is to maintain stable operation. Based on the general concept of homeostatic plasticity, we propose a simple compositional model of adaptive networks that achieve invariance with regard to undesired changes in the statistical properties of their input signals and maintain outputs with well-defined joint statistics. To achieve such invariance, the network model combines two functionally distinct types of plasticity. An abstract stochastic process neuron model implements a generalized form of intrinsic plasticity that adapts marginal statistics, relying only on mechanisms locally confined within each neuron and operating continuously in time, while a simple form of Hebbian synaptic plasticity operates on synaptic connections, thus shaping the interrelation between neurons as captured by a copula function. The combined effect of both mechanisms allows a neuron population to discover invariant representations of its inputs that remain stable under a wide range of transformations (e.g., shifting, scaling and (affine linear) mixing). The probabilistic model of homeostatic adaptation on a population level as presented here allows us to isolate and study the individual and the interaction dynamics of both mechanisms of plasticity and could guide the future search for computationally beneficial types of adaptation.

Cortical Spike Synchrony as a Measure of Input Familiarity.

Spike synchrony, which occurs in various cortical areas in response to specific perception, action, and memory tasks, has sparked a long-standing debate on the nature of temporal organization in cortex. One prominent view is that this type of synchrony facilitates the binding or grouping of separate stimulus components. We argue instead for a more general function: a measure of the prior probability of incoming stimuli, implemented by long-range, horizontal, intracortical connections. We show that networks of this kind-pulse-coupled excitatory spiking networks in a noisy environment-can provide a sufficient substrate for stimulus-dependent spike synchrony. This allows for a quick (few spikes) estimate of the match between inputs and the input history as encoded in the network structure. Given the ubiquity of small, strongly excitatory subnetworks in cortex, we thus propose that many experimental observations of spike synchrony can be viewed as signs of input patterns that resemble long-term experience-that is, of patterns with high prior probability.

Homeostatic plasticity for single node delay-coupled reservoir computing.

Supplementing a differential equation with delays results in an infinite-dimensional dynamical system. This property provides the basis for a reservoir computing architecture, where the recurrent neural network is replaced by a single nonlinear node, delay-coupled to itself. Instead of the spatial topology of a network, subunits in the delay-coupled reservoir are multiplexed in time along one delay span of the system. The computational power of the reservoir is contingent on this temporal multiplexing. Here, we learn optimal temporal multiplexing by means of a biologically inspired homeostatic plasticity mechanism. Plasticity acts locally and changes the distances between the subunits along the delay, depending on how responsive these subunits are to the input. After analytically deriving the learning mechanism, we illustrate its role in improving the reservoir's computational power. To this end, we investigate, first, the increase of the reservoir's memory capacity. Second, we predict a NARMA-10 time series, showing that plasticity reduces the normalized root-mean-square error by more than 20%. Third, we discuss plasticity's influence on the reservoir's input-information capacity, the coupling strength between subunits, and the distribution of the readout coefficients.

A Statistical Framework to Infer Delay and Direction of Information Flow from Measurements of Complex Systems.

In neuroscience, data are typically generated from neural network activity. The resulting time series represent measurements from spatially distributed subsystems with complex interactions, weakly coupled to a high-dimensional global system. We present a statistical framework to estimate the direction of information flow and its delay in measurements from systems of this type. Informed by differential topology, gaussian process regression is employed to reconstruct measurements of putative driving systems from measurements of the driven systems. These reconstructions serve to estimate the delay of the interaction by means of an analytical criterion developed for this purpose. The model accounts for a range of possible sources of uncertainty, including temporally evolving intrinsic noise, while assuming complex nonlinear dependencies. Furthermore, we show that if information flow is delayed, this approach also allows for inference in strong coupling scenarios of systems exhibiting synchronization phenomena. The validity of the method is demonstrated with a variety of delay-coupled chaotic oscillators. In addition, we show that these results seamlessly transfer to local field potentials in cat visual cortex.

Spatiotemporal computations of an excitable and plastic brain: neuronal plasticity leads to noise-robust and noise-constructive computations.

It is a long-established fact that neuronal plasticity occupies the central role in generating neural function and computation. Nevertheless, no unifying account exists of how neurons in a recurrent cortical network learn to compute on temporally and spatially extended stimuli. However, these stimuli constitute the norm, rather than the exception, of the brain's input. Here, we introduce a geometric theory of learning spatiotemporal computations through neuronal plasticity. To that end, we rigorously formulate the problem of neural representations as a relation in space between stimulus-induced neural activity and the asymptotic dynamics of excitable cortical networks. Backed up by computer simulations and numerical analysis, we show that two canonical and widely spread forms of neuronal plasticity, that is, spike-timing-dependent synaptic plasticity and intrinsic plasticity, are both necessary for creating neural representations, such that these computations become realizable. Interestingly, the effects of these forms of plasticity on the emerging neural code relate to properties necessary for both combating and utilizing noise. The neural dynamics also exhibits features of the most likely stimulus in the network's spontaneous activity. These properties of the spatiotemporal neural code resulting from plasticity, having their grounding in nature, further consolidate the biological relevance of our findings.

Spike synchrony as a measure of Gestalt structure.

The function of spike synchrony is debatable: some researchers view it as a mechanism for binding perceptual features, others - as a byproduct of brain activity. We argue for an alternative computational role: synchrony can estimate the prior probability of incoming stimuli. In V1, this can be achieved by comparing input with previously acquired visual experience, which is encoded in plastic horizontal intracortical connections. V1 connectivity structure can encode the acquired visual experience in the form of its aggregate statistics. Since the aggregate statistics of natural images tend to follow the Gestalt principles, we can assume that V1 is more often exposed to Gestalt-like stimuli, and this is manifested in its connectivity structure. At the same time, the connectivity structure has an impact on spike synchrony in V1. We used a spiking model with V1-like connectivity to demonstrate that spike synchrony reflects the Gestalt structure of the stimulus. We conducted simulation experiments with three Gestalt laws: proximity, similarity, and continuity, and found substantial differences in firing synchrony for stimuli with varying degrees of Gestalt-likeness. This allows us to conclude that spike synchrony indeed reflects the Gestalt structure of the stimulus, which can be interpreted as a mechanism for prior probability estimation.

Impact of spike train autostructure on probability distribution of joint spike events.

The discussion whether temporally coordinated spiking activity really exists and whether it is relevant has been heated over the past few years. To investigate this issue, several approaches have been taken to determine whether synchronized events occur significantly above chance, that is, whether they occur more often than expected if the neurons fire independently. Most investigations ignore or destroy the autostructure of the spiking activity of individual cells or assume Poissonian spiking as a model. Such methods that ignore the autostructure can significantly bias the coincidence statistics. Here, we study the influence of the autostructure on the probability distribution of coincident spiking events between tuples of mutually independent non-Poisson renewal processes. In particular, we consider two types of renewal processes that were suggested as appropriate models of experimental spike trains: a gamma and a log-normal process. For a gamma process, we characterize the shape of the distribution analytically with the Fano factor (FFc). In addition, we perform Monte Carlo estimations to derive the full shape of the distribution and the probability for false positives if a different process type is assumed as was actually present. We also determine how manipulations of such spike trains, here dithering, used for the generation of surrogate data change the distribution of coincident events and influence the significance estimation. We find, first, that the width of the coincidence count distribution and its FFc depend critically and in a nontrivial way on the detailed properties of the structure of the spike trains as characterized by the coefficient of variation CV. Second, the dependence of the FFc on the CV is complex and mostly nonmonotonic. Third, spike dithering, even if as small as a fraction of the interspike interval, can falsify the inference on coordinated firing.

Encoding through patterns: regression tree-based neuronal population models.

Although the existence of correlated spiking between neurons in a population is well known, the role such correlations play in encoding stimuli is not. We address this question by constructing pattern-based encoding models that describe how time-varying stimulus drive modulates the expression probabilities of population-wide spike patterns. The challenge is that large populations may express an astronomical number of unique patterns, and so fitting a unique encoding model for each individual pattern is not feasible. We avoid this combinatorial problem using a dimensionality-reduction approach based on regression trees. Using the insight that some patterns may, from the perspective of encoding, be statistically indistinguishable, the tree divisively clusters the observed patterns into groups whose member patterns possess similar encoding properties. These groups, corresponding to the leaves of the tree, are much smaller in number than the original patterns, and the tree itself constitutes a tractable encoding model for each pattern. Our formalism can detect an extremely weak stimulus-driven pattern structure and is based on maximizing the data likelihood, not making a priori assumptions as to how patterns should be grouped. Most important, by comparing pattern encodings with independent neuron encodings, one can determine if neurons in the population are driven independently or collectively. We demonstrate this method using multiple unit recordings from area 17 of anesthetized cat in response to a sinusoidal grating and show that pattern-based encodings are superior to those of independent neuron models. The agnostic nature of our clustering approach allows us to investigate encoding by the collective statistics that are actually present rather than those (such as pairwise) that might be presumed.

Mapping of visual receptive fields by tomographic reconstruction.

The moving bar experiment is a classic paradigm for characterizing the receptive field (RF) properties of neurons in primary visual cortex (V1). Current approaches for analyzing neural spiking activity recorded from these experiments do not take into account the point-process nature of these data and the circular geometry of the stimulus presentation. We present a novel analysis approach to mapping V1 receptive fields that combines point-process generalized linear models (PPGLM) with tomographic reconstruction computed by filtered-back projection. We use the method to map the RF sizes and orientations of 251 V1 neurons recorded from two macaque monkeys during a moving bar experiment. Our cross-validated goodness-of-fit analyses show that the PPGLM provides a more accurate characterization of spike train data than analyses based on rate functions computed by the methods of spike-triggered averages or first-order Wiener-Volterra kernel. Our analysis leads to a new definition of RF size as the spatial area over which the spiking activity is significantly greater than baseline activity. Our approach yields larger RF sizes and sharper orientation tuning estimates. The tomographic reconstruction paradigm further suggests an efficient approach to choosing the number of directions and the number of trials per direction in designing moving bar experiments. Our results demonstrate that standard tomographic principles for image reconstruction can be adapted to characterize V1 RFs and that two fundamental properties, size and orientation, may be substantially different from what is currently reported.

Transfer entropy--a model-free measure of effective connectivity for the neurosciences.

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.

Applying the multivariate time-rescaling theorem to neural population models.

Statistical models of neural activity are integral to modern neuroscience. Recently interest has grown in modeling the spiking activity of populations of simultaneously recorded neurons to study the effects of correlations and functional connectivity on neural information processing. However, any statistical model must be validated by an appropriate goodness-of-fit test. Kolmogorov-Smirnov tests based on the time-rescaling theorem have proven to be useful for evaluating point-process-based statistical models of single-neuron spike trains. Here we discuss the extension of the time-rescaling theorem to the multivariate (neural population) case. We show that even in the presence of strong correlations between spike trains, models that neglect couplings between neurons can be erroneously passed by the univariate time-rescaling test. We present the multivariate version of the time-rescaling theorem and provide a practical step-by-step procedure for applying it to testing the sufficiency of neural population models. Using several simple analytically tractable models and more complex simulated and real data sets, we demonstrate that important features of the population activity can be detected only using the multivariate extension of the test.

Dynamical relaying can yield zero time lag neuronal synchrony despite long conduction delays.

Multielectrode recordings have revealed zero time lag synchronization among remote cerebral cortical areas. However, the axonal conduction delays among such distant regions can amount to several tens of milliseconds. It is still unclear which mechanism is giving rise to isochronous discharge of widely distributed neurons, despite such latencies. Here, we investigate the synchronization properties of a simple network motif and found that, even in the presence of large axonal conduction delays, distant neuronal populations self-organize into lag-free oscillations. According to our results, cortico-cortical association fibers and certain cortico-thalamo-cortical loops represent ideal circuits to circumvent the phase shifts and time lags associated with conduction delays.

A trajectory-based loss function to learn missing terms in bifurcating dynamical systems.

Missing terms in dynamical systems are a challenging problem for modeling. Recent developments in the combination of machine learning and dynamical system theory open possibilities for a solution. We show how physics-informed differential equations and machine learning-combined in the Universal Differential Equation (UDE) framework by Rackauckas et al.-can be modified to discover missing terms in systems that undergo sudden fundamental changes in their dynamical behavior called bifurcations. With this we enable the application of the UDE approach to a wider class of problems which are common in many real world applications. The choice of the loss function, which compares the training data trajectory in state space and the current estimated solution trajectory of the UDE to optimize the solution, plays a crucial role within this approach. The Mean Square Error as loss function contains the risk of a reconstruction which completely misses the dynamical behavior of the training data. By contrast, our suggested trajectory-based loss function which optimizes two largely independent components, the length and angle of state space vectors of the training data, performs reliable well in examples of systems from neuroscience, chemistry and biology showing Saddle-Node, Pitchfork, Hopf and Period-doubling bifurcations.

Biologically Inspired Deep Learning Model for Efficient Foveal-Peripheral Vision.

While abundant in biology, foveated vision is nearly absent from computational models and especially deep learning architectures. Despite considerable hardware improvements, training deep neural networks still presents a challenge and constraints complexity of models. Here we propose an end-to-end neural model for foveal-peripheral vision, inspired by retino-cortical mapping in primates and humans. Our model has an efficient sampling technique for compressing the visual signal such that a small portion of the scene is perceived in high resolution while a large field of view is maintained in low resolution. An attention mechanism for performing "eye-movements" assists the agent in collecting detailed information incrementally from the observed scene. Our model achieves comparable results to a similar neural architecture trained on full-resolution data for image classification and outperforms it at video classification tasks. At the same time, because of the smaller size of its input, it can reduce computational effort tenfold and uses several times less memory. Moreover, we present an easy to implement bottom-up and top-down attention mechanism which relies on task-relevant features and is therefore a convenient byproduct of the main architecture. Apart from its computational efficiency, the presented work provides means for exploring active vision for agent training in simulated environments and anthropomorphic robotics.

Learning sparse and meaningful representations through embodiment.

How do humans acquire a meaningful understanding of the world with little to no supervision or semantic labels provided by the environment? Here we investigate embodiment with a closed loop between action and perception as one key component in this process. We take a close look at the representations learned by a deep reinforcement learning agent that is trained with high-dimensional visual observations collected in a 3D environment with very sparse rewards. We show that this agent learns stable representations of meaningful concepts such as doors without receiving any semantic labels. Our results show that the agent learns to represent the action relevant information, extracted from a simulated camera stream, in a wide variety of sparse activation patterns. The quality of the representations learned shows the strength of embodied learning and its advantages over fully supervised approaches.