Microtiming Deviations and Swing Feel in Jazz.

Jazz music that swings has the fascinating power to elicit a pleasant sensation of flow in listeners and the desire to synchronize body movements with the music. Whether microtiming deviations (MTDs), i.e. small timing deviations below the bar or phrase level, enhance the swing feel is highly debated in the current literature. Studies on other groove related genres did not find evidence for a positive impact of MTDs. The present study addresses jazz music and swing in particular, as there is some evidence that microtiming patterns are genre-specific. We recorded twelve piano jazz standards played by a professional pianist and manipulated the natural MTDs of the recordings in systematic ways by quantizing, expanding and inverting them. MTDs were defined with respect to a grid determined by the average swing ratio. The original and manipulated versions were presented in an online survey and evaluated by 160 listeners with various musical skill levels and backgrounds. Across pieces the quantized versions (without MTDs) were rated slightly higher and versions with expanded MTDs were rated lower with regard to swing than the original recordings. Unexpectedly, inversion had no impact on swing ratings except for two pieces. Our results suggest that naturally fluctuating MTDs are not an essential factor for the swing feel.

Correlated microtiming deviations in jazz and rock music.

Musical rhythms performed by humans typically show temporal fluctuations. While they have been characterized in simple rhythmic tasks, it is an open question what is the nature of temporal fluctuations, when several musicians perform music jointly in all its natural complexity. To study such fluctuations in over 100 original jazz and rock/pop recordings played with and without metronome we developed a semi-automated workflow allowing the extraction of cymbal beat onsets with millisecond precision. Analyzing the inter-beat interval (IBI) time series revealed evidence for two long-range correlated processes characterized by power laws in the IBI power spectral densities. One process dominates on short timescales (t < 8 beats) and reflects microtiming variability in the generation of single beats. The other dominates on longer timescales and reflects slow tempo variations. Whereas the latter did not show differences between musical genres (jazz vs. rock/pop), the process on short timescales showed higher variability for jazz recordings, indicating that jazz makes stronger use of microtiming fluctuations within a measure than rock/pop. Our results elucidate principles of rhythmic performance and can inspire algorithms for artificial music generation. By studying microtiming fluctuations in original music recordings, we bridge the gap between minimalistic tapping paradigms and expressive rhythmic performances.

Flexible information routing by transient synchrony.

Perception, cognition and behavior rely on flexible communication between microcircuits in distinct cortical regions. The mechanisms underlying rapid information rerouting between such microcircuits are still unknown. It has been proposed that changing patterns of coherence between local gamma rhythms support flexible information rerouting. The stochastic and transient nature of gamma oscillations in vivo, however, is hard to reconcile with such a function. Here we show that models of cortical circuits near the onset of oscillatory synchrony selectively route input signals despite the short duration of gamma bursts and the irregularity of neuronal firing. In canonical multiarea circuits, we find that gamma bursts spontaneously arise with matched timing and frequency and that they organize information flow by large-scale routing states. Specific self-organized routing states can be induced by minor modulations of background activity.

Model-Free Estimation of Tuning Curves and Their Attentional Modulation, Based on Sparse and Noisy Data.

Tuning curves are the functions that relate the responses of sensory neurons to various values within one continuous stimulus dimension (such as the orientation of a bar in the visual domain or the frequency of a tone in the auditory domain). They are commonly determined by fitting a model e.g. a Gaussian or other bell-shaped curves to the measured responses to a small subset of discrete stimuli in the relevant dimension. However, as neuronal responses are irregular and experimental measurements noisy, it is often difficult to determine reliably the appropriate model from the data. We illustrate this general problem by fitting diverse models to representative recordings from area MT in rhesus monkey visual cortex during multiple attentional tasks involving complex composite stimuli. We find that all models can be well-fitted, that the best model generally varies between neurons and that statistical comparisons between neuronal responses across different experimental conditions are affected quantitatively and qualitatively by specific model choices. As a robust alternative to an often arbitrary model selection, we introduce a model-free approach, in which features of interest are extracted directly from the measured response data without the need of fitting any model. In our attentional datasets, we demonstrate that data-driven methods provide descriptions of tuning curve features such as preferred stimulus direction or attentional gain modulations which are in agreement with fit-based approaches when a good fit exists. Furthermore, these methods naturally extend to the frequent cases of uncertain model selection. We show that model-free approaches can identify attentional modulation patterns, such as general alterations of the irregular shape of tuning curves, which cannot be captured by fitting stereotyped conventional models. Finally, by comparing datasets across different conditions, we demonstrate effects of attention that are cell- and even stimulus-specific. Based on these proofs-of-concept, we conclude that our data-driven methods can reliably extract relevant tuning information from neuronal recordings, including cells whose seemingly haphazard response curves defy conventional fitting approaches.

Paradoxical effects of coupling infectious livestock populations and imposing transport restrictions.

Spatial heterogeneity of a host population of mobile agents has been shown to be a crucial determinant of many aspects of disease dynamics, ranging from the proliferation of diseases to their persistence and to vaccination strategies. In addition, the importance of regional and structural differences grows in our modern world. Little is known, though, about the consequences when traits of a disease vary regionally. In this paper, we study the effect of a spatially varying per capita infection rate on the behaviour of livestock diseases. We show that the prevalence of an infectious livestock disease in a community of animals can paradoxically decrease owing to transport connections to other communities in which the risk of infection is higher. We study the consequences for the design of livestock transportation restriction measures and establish exact criteria to discriminate those connections that increase the level of infection in the community from those that decrease it.

Spike avalanches in vivo suggest a driven, slightly subcritical brain state.

In self-organized critical (SOC) systems avalanche size distributions follow power-laws. Power-laws have also been observed for neural activity, and so it has been proposed that SOC underlies brain organization as well. Surprisingly, for spiking activity in vivo, evidence for SOC is still lacking. Therefore, we analyzed highly parallel spike recordings from awake rats and monkeys, anesthetized cats, and also local field potentials from humans. We compared these to spiking activity from two established critical models: the Bak-Tang-Wiesenfeld model, and a stochastic branching model. We found fundamental differences between the neural and the model activity. These differences could be overcome for both models through a combination of three modifications: (1) subsampling, (2) increasing the input to the model (this way eliminating the separation of time scales, which is fundamental to SOC and its avalanche definition), and (3) making the model slightly sub-critical. The match between the neural activity and the modified models held not only for the classical avalanche size distributions and estimated branching parameters, but also for two novel measures (mean avalanche size, and frequency of single spikes), and for the dependence of all these measures on the temporal bin size. Our results suggest that neural activity in vivo shows a mélange of avalanches, and not temporally separated ones, and that their global activity propagation can be approximated by the principle that one spike on average triggers a little less than one spike in the next step. This implies that neural activity does not reflect a SOC state but a slightly sub-critical regime without a separation of time scales. Potential advantages of this regime may be faster information processing, and a safety margin from super-criticality, which has been linked to epilepsy.

Transfer entropy reconstruction and labeling of neuronal connections from simulated calcium imaging.

Neuronal dynamics are fundamentally constrained by the underlying structural network architecture, yet much of the details of this synaptic connectivity are still unknown even in neuronal cultures in vitro. Here we extend a previous approach based on information theory, the Generalized Transfer Entropy, to the reconstruction of connectivity of simulated neuronal networks of both excitatory and inhibitory neurons. We show that, due to the model-free nature of the developed measure, both kinds of connections can be reliably inferred if the average firing rate between synchronous burst events exceeds a small minimum frequency. Furthermore, we suggest, based on systematic simulations, that even lower spontaneous inter-burst rates could be raised to meet the requirements of our reconstruction algorithm by applying a weak spatially homogeneous stimulation to the entire network. By combining multiple recordings of the same in silico network before and after pharmacologically blocking inhibitory synaptic transmission, we show then how it becomes possible to infer with high confidence the excitatory or inhibitory nature of each individual neuron.

Possible origin of stagnation and variability of earth's biodiversity.

The magnitude and variability of Earth's biodiversity have puzzled scientists ever since paleontologic fossil databases became available. We identify and study a model of interdependent species where both endogenous and exogenous impacts determine the nonstationary extinction dynamics. The framework provides an explanation for the qualitative difference of marine and continental biodiversity growth. In particular, the stagnation of marine biodiversity may result from a global transition from an imbalanced to a balanced state of the species dependency network. The predictions of our framework are in agreement with paleontologic databases.

Optogenetic stimulation effectively enhances intrinsically generated network synchrony.

Synchronized bursting is found in many brain areas and has also been implicated in the pathophysiology of neuropsychiatric disorders such as epilepsy, Parkinson's disease, and schizophrenia. Despite extensive studies of network burst synchronization, it is insufficiently understood how this type of network wide synchronization can be strengthened, reduced, or even abolished. We combined electrical recording using multi-electrode array with optical stimulation of cultured channelrhodopsin-2 transducted hippocampal neurons to study and manipulate network burst synchronization. We found low frequency photo-stimulation protocols that are sufficient to induce potentiation of network bursting, modifying bursting dynamics, and increasing interneuronal synchronization. Surprisingly, slowly fading-in light stimulation, which substantially delayed and reduced light-driven spiking, was at least as effective in reorganizing network dynamics as much stronger pulsed light stimulation. Our study shows that mild stimulation protocols that do not enforce particular activity patterns onto the network can be highly effective inducers of network-level plasticity.

Compositionality in neural control: an interdisciplinary study of scribbling movements in primates.

This article discusses the compositional structure of hand movements by analyzing and modeling neural and behavioral data obtained from experiments where a monkey (Macaca fascicularis) performed scribbling movements induced by a search task. Using geometrically based approaches to movement segmentation, it is shown that the hand trajectories are composed of elementary segments that are primarily parabolic in shape. The segments could be categorized into a small number of classes on the basis of decreasing intra-class variance over the course of training. A separate classification of the neural data employing a hidden Markov model showed a coincidence of the neural states with the behavioral categories. An additional analysis of both types of data by a data mining method provided evidence that the neural activity patterns underlying the behavioral primitives were formed by sets of specific and precise spike patterns. A geometric description of the movement trajectories, together with precise neural timing data indicates a compositional variant of a realistic synfire chain model. This model reproduces the typical shapes and temporal properties of the trajectories; hence the structure and composition of the primitives may reflect meaningful behavior.

Intensity fluctuations of waves in random media: what is the semiclassical limit?

Waves traveling through weakly random media are known to be strongly affected by their corresponding ray dynamics, in particular in forming linear freak waves. The ray intensity distribution, which, e.g., quantifies the probability of freak waves is unknown, however, and a theory of how it is approached in an appropriate semiclassical limit of wave mechanics is lacking. We show that this limit is not the usual limit of small wavelengths, but that of decoherence. Our theory, which can describe the intensity distribution for an arbitrary degree of coherence is relevant to a wide range of physical systems, as decoherence is omnipresent in real systems.

Critical dynamics in associative memory networks.

Critical behavior in neural networks is characterized by scale-free avalanche size distributions and can be explained by self-regulatory mechanisms. Theoretical and experimental evidence indicates that information storage capacity reaches its maximum in the critical regime. We study the effect of structural connectivity formed by Hebbian learning on the criticality of network dynamics. The network only endowed with Hebbian learning does not allow for simultaneous information storage and criticality. However, the critical regime can be stabilized by short-term synaptic dynamics in the form of synaptic depression and facilitation or, alternatively, by homeostatic adaptation of the synaptic weights. We show that a heterogeneous distribution of maximal synaptic strengths does not preclude criticality if the Hebbian learning is alternated with periods of critical dynamics recovery. We discuss the relevance of these findings for the flexibility of memory in aging and with respect to the recent theory of synaptic plasticity.

Model-free reconstruction of excitatory neuronal connectivity from calcium imaging signals.

A systematic assessment of global neural network connectivity through direct electrophysiological assays has remained technically infeasible, even in simpler systems like dissociated neuronal cultures. We introduce an improved algorithmic approach based on Transfer Entropy to reconstruct structural connectivity from network activity monitored through calcium imaging. We focus in this study on the inference of excitatory synaptic links. Based on information theory, our method requires no prior assumptions on the statistics of neuronal firing and neuronal connections. The performance of our algorithm is benchmarked on surrogate time series of calcium fluorescence generated by the simulated dynamics of a network with known ground-truth topology. We find that the functional network topology revealed by Transfer Entropy depends qualitatively on the time-dependent dynamic state of the network (bursting or non-bursting). Thus by conditioning with respect to the global mean activity, we improve the performance of our method. This allows us to focus the analysis to specific dynamical regimes of the network in which the inferred functional connectivity is shaped by monosynaptic excitatory connections, rather than by collective synchrony. Our method can discriminate between actual causal influences between neurons and spurious non-causal correlations due to light scattering artifacts, which inherently affect the quality of fluorescence imaging. Compared to other reconstruction strategies such as cross-correlation or Granger Causality methods, our method based on improved Transfer Entropy is remarkably more accurate. In particular, it provides a good estimation of the excitatory network clustering coefficient, allowing for discrimination between weakly and strongly clustered topologies. Finally, we demonstrate the applicability of our method to analyses of real recordings of in vitro disinhibited cortical cultures where we suggest that excitatory connections are characterized by an elevated level of clustering compared to a random graph (although not extreme) and can be markedly non-local.

Dynamic effective connectivity of inter-areal brain circuits.

Anatomic connections between brain areas affect information flow between neuronal circuits and the synchronization of neuronal activity. However, such structural connectivity does not coincide with effective connectivity (or, more precisely, causal connectivity), related to the elusive question "Which areas cause the present activity of which others?". Effective connectivity is directed and depends flexibly on contexts and tasks. Here we show that dynamic effective connectivity can emerge from transitions in the collective organization of coherent neural activity. Integrating simulation and semi-analytic approaches, we study mesoscale network motifs of interacting cortical areas, modeled as large random networks of spiking neurons or as simple rate units. Through a causal analysis of time-series of model neural activity, we show that different dynamical states generated by a same structural connectivity motif correspond to distinct effective connectivity motifs. Such effective motifs can display a dominant directionality, due to spontaneous symmetry breaking and effective entrainment between local brain rhythms, although all connections in the considered structural motifs are reciprocal. We show then that transitions between effective connectivity configurations (like, for instance, reversal in the direction of inter-areal interactions) can be triggered reliably by brief perturbation inputs, properly timed with respect to an ongoing local oscillation, without the need for plastic synaptic changes. Finally, we analyze how the information encoded in spiking patterns of a local neuronal population is propagated across a fixed structural connectivity motif, demonstrating that changes in the active effective connectivity regulate both the efficiency and the directionality of information transfer. Previous studies stressed the role played by coherent oscillations in establishing efficient communication between distant areas. Going beyond these early proposals, we advance here that dynamic interactions between brain rhythms provide as well the basis for the self-organized control of this "communication-through-coherence", making thus possible a fast "on-demand" reconfiguration of global information routing modalities.

The nature and perception of fluctuations in human musical rhythms.

Although human musical performances represent one of the most valuable achievements of mankind, the best musicians perform imperfectly. Musical rhythms are not entirely accurate and thus inevitably deviate from the ideal beat pattern. Nevertheless, computer generated perfect beat patterns are frequently devalued by listeners due to a perceived lack of human touch. Professional audio editing software therefore offers a humanizing feature which artificially generates rhythmic fluctuations. However, the built-in humanizing units are essentially random number generators producing only simple uncorrelated fluctuations. Here, for the first time, we establish long-range fluctuations as an inevitable natural companion of both simple and complex human rhythmic performances. Moreover, we demonstrate that listeners strongly prefer long-range correlated fluctuations in musical rhythms. Thus, the favorable fluctuation type for humanizing interbeat intervals coincides with the one generically inherent in human musical performances.

Heat transport in active harmonic chains.

We show that a harmonic lattice model with amplifying and attenuating elements, when coupled to two thermal baths, exhibits unique heat transport properties. Some of these novel features include anomalous nonequilibrium steady-state heat currents, negative differential thermal conductance, as well as nonreciprocal heat transport. We find that when these elements are arranged in a PT-symmetric manner, the domain of existence of the nonequilibrium steady state is maximized. We propose an electronic experimental setup based on resistive-inductive-capacitive (RLC) transmission lines, where our predictions can be tested.

Spike correlations - what can they tell about synchrony?

Sensory and cognitive processing relies on the concerted activity of large populations of neurons. The advent of modern experimental techniques like two-photon population calcium imaging makes it possible to monitor the spiking activity of multiple neurons as they are participating in specific cognitive tasks. The development of appropriate theoretical tools to quantify and interpret the spiking activity of multiple neurons, however, is still in its infancy. One of the simplest and widely used measures of correlated activity is the pairwise correlation coefficient. While spike correlation coefficients are easy to compute using the available numerical toolboxes, it has remained largely an open question whether they are indeed a reliable measure of synchrony. Surprisingly, despite the intense use of correlation coefficients in the design of synthetic spike trains, the construction of population models and the assessment of the synchrony level in live neuronal networks very little was known about their computational properties. We showed that many features of pairwise spike correlations can be studied analytically in a tractable threshold model. Importantly, we demonstrated that under some circumstances the correlation coefficients can vanish, even though input and also pairwise spike cross correlations are present. This finding suggests that the most popular and frequently used measures can, by design, fail to capture the neuronal synchrony.

Universal statistics of branched flows.

Even very weak correlated disorder potentials can cause extreme fluctuations in Hamiltonian flows. In two dimensions this leads to a pronounced branching of the flow. Although present in a great variety of physical systems, a quantitative theory of the branching statistics is lacking. Here, we derive an analytical expression for the number of branches valid for all distances from a source. We also derive the scaling relations that make this expression universal for a wide range of random potentials. Our theory has possible applications in many fields ranging from semiconductor to geophysics.

Signatures of synchrony in pairwise count correlations.

Concerted neural activity can reflect specific features of sensory stimuli or behavioral tasks. Correlation coefficients and count correlations are frequently used to measure correlations between neurons, design synthetic spike trains and build population models. But are correlation coefficients always a reliable measure of input correlations? Here, we consider a stochastic model for the generation of correlated spike sequences which replicate neuronal pairwise correlations in many important aspects. We investigate under which conditions the correlation coefficients reflect the degree of input synchrony and when they can be used to build population models. We find that correlation coefficients can be a poor indicator of input synchrony for some cases of input correlations. In particular, count correlations computed for large time bins can vanish despite the presence of input correlations. These findings suggest that network models or potential coding schemes of neural population activity need to incorporate temporal properties of correlated inputs and take into consideration the regimes of firing rates and correlation strengths to ensure that their building blocks are an unambiguous measures of synchrony.

Correlations and synchrony in threshold neuron models.

We study how threshold models and neocortical neurons transfer temporal and interneuronal input correlations to correlations of spikes. In both, we find that the low common input regime is governed by firing rate dependent spike correlations which are sensitive to the detailed structure of input correlation functions. In the high common input regime, the spike correlations are largely insensitive to the firing rate and exhibit a universal peak shape. We further show that pairs with different firing rates driven by common inputs in general exhibit asymmetric spike correlations.