Geometric Analysis of Synchronization in Neuronal Networks with Global Inhibition and Coupling Delays

We study synaptically coupled neuronal networks to identify the role of coupling delays in network's synchronized behaviors. We consider a network of excitable, relaxation oscillator neurons where two distinct populations, one excitatory and one inhibitory, are coupled and interact with each other. The excitatory population is uncoupled, while the inhibitory population is tightly coupled. A geometric singular perturbation analysis yields existence and stability conditions for synchronization states under different firing patterns between the two populations, along with formulas for the periods of such synchronous solutions. Our results demonstrate that the presence of coupling delays in the network promotes synchronization. Numerical simulations are conducted to supplement and validate analytical results. We show the results carry over to a model for spindle sleep rhythms in thalamocortical networks, one of the biological systems which motivated our study. The analysis helps to explain how coupling delays in either excitatory or inhibitory synapses contribute to producing synchronized rhythms.Comment: 43 pages, 12 figure

Bio-inspired Dynamic Control Systems with Time Delays

The world around us exhibits a rich and ever changing environment of startling, bewildering and fascinating complexity. Almost everything is never as simple as it seems, but through the chaos we may catch fleeting glimpses of the mechanisms within. Throughout the history of human endeavour we have mimicked nature to harness it for our own ends. Our attempts to develop truly autonomous and intelligent machines have however struggled with the limitations of our human ability. This has encouraged some to shirk this responsibility and instead model biological processes and systems to do it for us. This Thesis explores the introduction of continuous time delays into biologically inspired dynamic control systems. We seek to exploit rich temporal dynamics found in physical and biological systems for modelling complex or adaptive behaviour through the artificial evolution of networks to control robots. Throughout, arguments have been presented for the modelling of delays not only to better represent key facets of physical and biological systems, but to increase the computational potential of such systems for the synthesis of control. The thorough investigation of the dynamics of small delayed networks with a wide range of time delays has been undertaken, with a detailed mathematical description of the fixed points of the system and possible oscillatory modes developed to fully describe the behaviour of a single node. Exploration of the behaviour for even small delayed networks illustrates the range of complex behaviour possible and guides the development of interesting solutions. To further exploit the potential of the rich dynamics in such systems, a novel approach to the 3D simulation of locomotory robots has been developed focussing on minimising the computational cost. To verify this simulation tool a simple quadruped robot was developed and the motion of the robot when undergoing a manually designed gait evaluated. The results displayed a high degree of agreement between the simulation and laser tracker data, verifying the accuracy of the model developed. A new model of a dynamic system which includes continuous time delays has been introduced, and its utility demonstrated in the evolution of networks for the solution of simple learning behaviours. A range of methods has been developed for determining the time delays, including the novel concept of representing the time delays as related to the distance between nodes in a spatial representation of the network. The application of these tools to a range of examples has been explored, from Gene Regulatory Networks (GRNs) to robot control and neural networks. The performance of these systems has been compared and contrasted with the efficacy of evolutionary runs for the same task over the whole range of network and delay types. It has been shown that delayed dynamic neural systems are at least as capable as traditional Continuous Time Recurrent Neural Networks (CTRNNs) and show significant performance improvements in the control of robot gaits. Experiments in adaptive behaviour, where there is not such a direct link between the enhanced system dynamics and performance, showed no such discernible improvement. Whilst we hypothesise that the ability of such delayed networks to generate switched pattern generating nodes may be useful in Evolutionary Robotics (ER) this was not borne out here. The spatial representation of delays was shown to be more efficient for larger networks, however these techniques restricted the search to lower complexity solutions or led to a significant falloff as the network structure becomes more complex. This would suggest that for anything other than a simple genotype, the direct method for encoding delays is likely most appropriate. With proven benefits for robot locomotion and the open potential for adaptive behaviour delayed dynamic systems for evolved control remain an interesting and promising field in complex systems research

Dual coding with STDP in a spiking recurrent neural network model of the hippocampus.

The firing rate of single neurons in the mammalian hippocampus has been demonstrated to encode for a range of spatial and non-spatial stimuli. It has also been demonstrated that phase of firing, with respect to the theta oscillation that dominates the hippocampal EEG during stereotype learning behaviour, correlates with an animal's spatial location. These findings have led to the hypothesis that the hippocampus operates using a dual (rate and temporal) coding system. To investigate the phenomenon of dual coding in the hippocampus, we examine a spiking recurrent network model with theta coded neural dynamics and an STDP rule that mediates rate-coded Hebbian learning when pre- and post-synaptic firing is stochastic. We demonstrate that this plasticity rule can generate both symmetric and asymmetric connections between neurons that fire at concurrent or successive theta phase, respectively, and subsequently produce both pattern completion and sequence prediction from partial cues. This unifies previously disparate auto- and hetero-associative network models of hippocampal function and provides them with a firmer basis in modern neurobiology. Furthermore, the encoding and reactivation of activity in mutually exciting Hebbian cell assemblies demonstrated here is believed to represent a fundamental mechanism of cognitive processing in the brain

Spatially structured oscillations in a two-dimensional excitatory neuronal network with synaptic depression

We study the spatiotemporal dynamics of a two-dimensional excitatory neuronal network with synaptic depression. Coupling between populations of neurons is taken to be nonlocal, while depression is taken to be local and presynaptic. We show that the network supports a wide range of spatially structured oscillations, which are suggestive of phenomena seen in cortical slice experiments and in vivo. The particular form of the oscillations depends on initial conditions and the level of background noise. Given an initial, spatially localized stimulus, activity evolves to a spatially localized oscillating core that periodically emits target waves. Low levels of noise can spontaneously generate several pockets of oscillatory activity that interact via their target patterns. Periodic activity in space can also organize into spiral waves, provided that there is some source of rotational symmetry breaking due to external stimuli or noise. In the high gain limit, no oscillatory behavior exists, but a transient stimulus can lead to a single, outward propagating target wave

A Scalable Model of Cerebellar Adaptive Timing and Sequencing: The Recurrent Slide and Latch (RSL) Model

From the dawn of modern neural network theory, the mammalian cerebellum has been a favored object of mathematical modeling studies. Early studies focused on the fan-out, convergence, thresholding, and learned weighting of perceptual-motor signals within the cerebellar cortex. This led in the proposals of Albus (1971; 1975) and Marr (1969) to the still viable idea that the granule cell stage in the cerebellar cortex performs a sparse expansive recoding of the time-varying input vector. This recoding reveals and emphasizes combinations (of input state variables) in a distributed representation that serves as a basis for the learned, state-dependent control actions engendered by cerebellar outputs to movement related centers. Although well-grounded as such, this perspective seriously underestimates the intelligence of the cerebellar cortex. Context and state information arises asynchronously due to the heterogeneity of sources that contribute signals to compose the cerebellar input vector. These sources include radically different sensory systems - vision, kinesthesia, touch, balance and audition - as well as many stages of the motor output channel. To make optimal use of available signals, the cerebellum must be able to sift the evolving state representation for the most reliable predictors of the need for control actions, and to use those predictors even if they appear only transiently and well in advance of the optimal time for initiating the control action. Such a cerebellar adaptive timing competence has recently been experimentally verified (Perrett, Ruiz, & Mauk, 1993). This paper proposes a modification to prior, population, models for cerebellar adaptive timing and sequencing. Since it replaces a population with a single clement, the proposed Recurrent Slide and Latch (RSL) model is in one sense maximally efficient, and therefore optimal from the perspective of scalability.Defense Advanced Research Projects Agency and the Office of Naval Research (N00014-92-J-1309, N00014-93-1-1364, N00014-95-1-0409)

Bio-inspired vision-based leader-follower formation flying in the presence of delays

Flocking starlings at dusk are known for the mesmerizing and intricate shapes they generate, as well as how fluid these shapes change. They seem to do this effortlessly. Real-life vision-based flocking has not been achieved in micro-UAVs (micro Unmanned Aerial Vehicles) to date. Towards this goal, we make three contributions in this paper: (i) we used a computational approach to develop a bio-inspired architecture for vision-based Leader-Follower formation flying on two micro-UAVs. We believe that the minimal computational cost of the resulting algorithm makes it suitable for object detection and tracking during high-speed flocking; (ii) we show that provided delays in the control loop of a micro-UAV are below a critical value, Kalman filter-based estimation algorithms are not required to achieve Leader-Follower formation flying; (iii) unlike previous approaches, we do not use external observers, such as GPS signals or synchronized communication with flock members. These three contributions could be useful in achieving vision-based flocking in GPS-denied environments on computationally-limited agents

Neural models of learning and visual grouping in the presence of finite conduction velocities

The hypothesis of object binding-by-synchronization in the visual cortex has been supported by recent experiments in awake monkeys. They demonstrated coherence among gamma-activities (30–90 Hz) of local neural groups and its perceptual modulation according to the rules of figure-ground segregation. Interactions within and between these neural groups are based on axonal spike conduction with finite velocities. Physiological studies confirmed that the majority of transmission delays is comparable to the temporal scale defined by gamma-activity (11–33 ms). How do these finite velocities influence the development of synaptic connections within and between visual areas? What is the relationship between the range of gamma-coherence and the velocity of signal transmission? Are these large temporal delays compatible with recently discovered phenomenon of gamma-waves traveling across larger parts of the primary visual cortex? The refinement of connections in the immature visual cortex depends on temporal Hebbian learning to adjust synaptic efficacies between spiking neurons. The impact of constant, finite, axonal spike conduction velocities on this process was investigated using a set of topographic network models. Random spike trains with a confined temporal correlation width mimicked cortical activity before visual experience. After learning, the lateral connectivity within one network layer became spatially restricted, the width of the connection profile being directly proportional to the lateral conduction velocity. Furthermore, restricted feedforward divergence developed between neurons of two successive layers. The size of this connection profile matched the lateral connection profile of the lower layer neuron. The mechanism in this network model is suitable to explain the emergence of larger receptive fields at higher visual areas while preserving a retinotopic mapping. The influence of finite conduction velocities on the local generation of gamma-activities and their spatial synchronization was investigated in a model of a mature visual area. Sustained input and local inhibitory feedback was sufficient for the emergence of coherent gamma-activity that extended across few millimeters. Conduction velocities had a direct impact on the frequency of gamma-oscillations, but did neither affect gamma-power nor the spatial extent of gamma-coherence. Adding long-range horizontal connections between excitatory neurons, as found in layer 2/3 of the primary visual cortex, increased the spatial range of gamma-coherence. The range was maximal for zero transmission delays, and for all distances attenuated with finite, decreasing lateral conduction velocities. Below a velocity of 0.5 m/s, gamma-power and gamma-coherence were even smaller than without these connections at all, i.e., slow horizontal connections actively desynchronized neural populations. In conclusion, the enhancement of gamma-coherence by horizontal excitatory connections critically depends on fast conduction velocities. Coherent gamma-activity in the primary visual cortex and the accompanying models was found to only cover small regions of the visual field. This challenges the role of gamma-synchronization to solve the binding problem for larger object representations. Further analysis of the previous model revealed that the patches of coherent gamma-activity (1.8 mm half-height decline) were part of more globally occurring gamma-waves, which coupled over much larger distances (6.3 mm half-height decline). The model gamma-waves observed here are very similar to those found in the primary visual cortex of awake monkeys, indicating that local recurrent inhibition and restricted horizontal connections with finite axonal velocities are sufficient requirements for their emergence. In conclusion, since the model is in accordance with the connectivity and gamma-processes in the primary visual cortex, the results support the hypothesis that gamma-waves provide a generalized concept for object binding in the visual cortex