Control of movement of underwater swimmers: Animals, simulated animates and swimming robots.

The control of movement in living organisms represents a fundamental task that the brain has evolved to solve. One crucial aspect is how the nervous system organizes the transformation of sensory information into motor commands. These commands lead to muscle activation and subsequent animal movement, which can exhibit complex patterns. One example of such movement is locomotion, which involves the translation of the entire body through space. Central Pattern Generators (CPGs) are neuronal circuits that provide control signals for these movements. Compared to the intricate circuits found in the brain, CPGs can be simplified into networks of neurons that generate rhythmic activation, coordinating muscle movements. Since the 1990s, researchers have developed numerous models of locomotive circuits to simulate different types of animal movement, including walking, flying, and swimming. Initially, the primary goal of these studies was to construct biomimetic robots. However, it became apparent that simplified CPGs alone were not sufficient to replicate the diverse range of adaptive locomotive movements observed in living organisms. Factors such as sensory modulation, higher-level control, and cognitive components related to learning and memory needed to be considered. This necessitated the use of more complex, high-dimensional circuits, as well as novel materials and hardware, in both modeling and robotics. With advancements in high-power computing, artificial intelligence, big data processing, smart materials, and electronics, the possibility of designing a new generation of true bio-mimetic robots has emerged. These robots have the capability to imitate not only simple locomotion but also exhibit adaptive motor behavior and decision-making. This motivation serves as the foundation for the current review, which aims to analyze existing concepts and models of movement control systems. As an illustrative example, we focus on underwater movement and explore the fundamental biological concepts, as well as the mathematical and physical models that underlie locomotion and its various modulations.

Tonic regulation of stationary asynchronous firing of a neural network.

The impact of tonic conductance upon population activity was investigated. An extra tonic transmembrane current through GABA-activated extrasynaptic GABA A -receptors was found to control stationary asynchronous firing both quantitatively and qualitatively. Quantitative regulation consisted in alterating a current level of stationary population activity while qualitative regulation manifested itself in appearance of resilient asynchronous spiking in case GABA reversal potential exceeded a certain threshold. The study was based on a modified rate model after Wilson and Cowan and backed up with a computer simulation of an explicit network model.

Oscillations in a neurite growth model with extracellular feedback.

We consider the influence of extracellular signalling on neurite elongation in a model of neurite growth mediated by building proteins (e.g., tubulin). The tubulin production dynamics were supplied by a function describing the influence of extracellular signalling, which can promote or depress neurite elongation. We found that this extracellular feedback could generate neurite length oscillations consisting of a periodic sequence of elongations and retractions. The oscillations prevent further outgrowth of the neurite, which becomes trapped in the non-uniform extracellular field. We analysed the characteristics of the elongation process for different distributions of attracting and repelling sources of the extracellular signalling molecules. The model predicts three different scenarios of neurite development in the extracellular field, including monotonic and oscillatory outgrowth, localised limit cycle oscillations and complete growth depression.

Bi-directional astrocytic regulation of neuronal activity within a network.

The concept of a tripartite synapse holds that astrocytes can affect both the pre- and post-synaptic compartments through the Ca(2+)-dependent release of gliotransmitters. Because astrocytic Ca(2+) transients usually last for a few seconds, we assumed that astrocytic regulation of synaptic transmission may also occur on the scale of seconds. Here, we considered the basic physiological functions of tripartite synapses and investigated astrocytic regulation at the level of neural network activity. The firing dynamics of individual neurons in a spontaneous firing network was described by the Hodgkin-Huxley model. The neurons received excitatory synaptic input driven by the Poisson spike train with variable frequency. The mean field concentration of the released neurotransmitter was used to describe the presynaptic dynamics. The amplitudes of the excitatory postsynaptic currents (PSCs) obeyed the gamma distribution law. In our model, astrocytes depressed the presynaptic release and enhanced the PSCs. As a result, low frequency synaptic input was suppressed while high frequency input was amplified. The analysis of the neuron spiking frequency as an indicator of network activity revealed that tripartite synaptic transmission dramatically changed the local network operation compared to bipartite synapses. Specifically, the astrocytes supported homeostatic regulation of the network activity by increasing or decreasing firing of the neurons. Thus, the astrocyte activation may modulate a transition of neural network into bistable regime of activity with two stable firing levels and spontaneous transitions between them.

Bistability induces episodic spike communication by inhibitory neurons in neuronal networks.

Bistability is one of the important features of nonlinear dynamical systems. In neurodynamics, bistability has been found in basic Hodgkin-Huxley equations describing the cell membrane dynamics. When the neuron is clamped near its threshold, the stable rest potential may coexist with the stable limit cycle describing periodic spiking. However, this effect is often neglected in network computations where the neurons are typically reduced to threshold firing units (e.g., integrate-and-fire models). We found that the bistability may induce spike communication by inhibitory coupled neurons in the spiking network. The communication is realized in the form of episodic discharges with synchronous (correlated) spikes during the episodes. A spiking phase map is constructed to describe the synchronization and to estimate basic spike phase locking modes.

Bifurcation mechanisms of regular and chaotic network signaling in brain astrocytes.

Bifurcation mechanisms underlying calcium oscillations in the network of astrocytes are investigated. Network model includes the dynamics of intracellular calcium concentration and intercellular diffusion of inositol 1,4,5-trisphosphate through gap junctions. Bifurcation analysis of underlying nonlinear dynamical system is presented. Parameter regions and principle bifurcation boundaries have been delineated and described. We show how variations of the diffusion rate can lead to generation of network calcium oscillations in originally nonoscillating cells. Different scenarios of regular activity and its transitions to chaotic dynamics have been obtained. Then, the bifurcations have been associated with statistical characteristics of calcium signals showing that different bifurcation scenarios yield qualitative changes in experimentally measurable quantities of the astrocyte activity, e.g., statistics of calcium spikes.

[Cluster associative memory formation in a three-layer network of phase oscillators].

A three-layer network model of oscillatory associative mermory is proposed. The network is capable to store binary images that can be retrieved if an appropriate stimulus has been applied. Binary images are encoded in the form of the spatial distribution of oscillatory phase clusters in-phase (+1) and anti-phase relative to the base periodic signal. The information is loaded into the network using a set of interlayer connection weights. A condition for error-free pattern retrieval has been obtained, which imposes a certain limitation on the maximal number of patterns to be stored in the memory (storage capacity). It has been shown that the capacity can be significantly increased by the generation of optimal pattern alphabet (basic pattern set). The number of stored patterns can reach values of the network size (the number of oscillators in the layer), which is significantly higher than the capacity of traditional oscillatory memory models. The dynamical and information characteristics of the retrieval process based on the optimal alphabet including the estimations of attraction basins and the admissible input pattern discrepancy for error-free retrieval have been investigated.

Spontaneous calcium signals induced by gap junctions in a network model of astrocytes.

The dynamics of a network model of astrocytes coupled by gap junctions is investigated. Calcium dynamics of the single cell is described by the biophysical model comprising the set of three nonlinear differential equations. Intercellular dynamics is provided by the diffusion of inositol 1,4,5-trisphosphate (IP3) through gap junctions between neighboring astrocytes. It is found that the diffusion induces the appearance of spontaneous activity patterns in the network. Stability of the network steady state is analyzed. It is proved that the increase of the diffusion coefficient above a certain critical value yields the generation of low-amplitude subthreshold oscillatory signals in a certain frequency range. It is shown that such spontaneous oscillations can facilitate calcium pulse generation and provide a certain time scale in astrocyte signaling.

Spiking dynamics of interacting oscillatory neurons.

Spiking sequences emerging from dynamical interaction in a pair of oscillatory neurons are investigated theoretically and experimentally. The model comprises two unidirectionally coupled FitzHugh-Nagumo units with modified excitability (MFHN). The first (master) unit exhibits a periodic spike sequence with a certain frequency. The second (slave) unit is in its excitable mode and responds on the input signal with a complex (chaotic) spike trains. We analyze the dynamic mechanisms underlying different response behavior depending on interaction strength. Spiking phase maps describing the response dynamics are obtained. Complex phase locking and chaotic sequences are investigated. We show how the response spike trains can be effectively controlled by the interaction parameter and discuss the problem of neuronal information encoding.

Self-referential phase reset based on inferior olive oscillator dynamics.

The olivo-cerebellar network is a key neuronal circuit that provides high-level motor control in the vertebrate CNS. Functionally, its network dynamics is organized around the oscillatory membrane potential properties of inferior olive (IO) neurons and their electrotonic connectivity. Because IO action potentials are generated at the peaks of the quasisinusoidal membrane potential oscillations, their temporal firing properties are defined by the IO rhythmicity. Excitatory inputs to these neurons can produce oscillatory phase shifts without modifying the amplitude or frequency of the oscillations, allowing well defined time-shift modulation of action potential generation. Moreover, the resulting phase is defined only by the amplitude and duration of the reset stimulus and is independent of the original oscillatory phase when the stimulus was delivered. This reset property, henceforth referred to as selfreferential phase reset, results in the generation of organized clusters of electrically coupled cells that oscillate in phase and are controlled by inhibitory feedback loops through the cerebellar nuclei and the cerebellar cortex. These clusters provide a dynamical representation of arbitrary motor intention patterns that are further mapped to the motor execution system. Being supplied with sensory inputs, the olivo-cerebellar network is capable of rearranging the clusters during the process of movement execution. Accordingly, the phase of the IO oscillators can be rapidly reset to a desired phase independently of the history of phase evolution. The goal of this article is to show how this selfreferential phase reset may be implemented into a motor control system by using a biologically based mathematical model.

Emergence of chaotic attractor and anti-synchronization for two coupled monostable neurons.

The dynamics of two coupled piece-wise linear one-dimensional monostable maps is investigated. The single map is associated with Poincare section of the FitzHugh-Nagumo neuron model. It is found that a diffusive coupling leads to the appearance of chaotic attractor. The attractor exists in an invariant region of phase space bounded by the manifolds of the saddle fixed point and the saddle periodic point. The oscillations from the chaotic attractor have a spike-burst shape with anti-phase synchronized spiking.

Olivo-cerebellar cluster-based universal control system.

The olivo-cerebellar network plays a key role in the organization of vertebrate motor control. The oscillatory properties of inferior olive (IO) neurons have been shown to provide timing signals for motor coordination in which spatio-temporal coherent oscillatory neuronal clusters control movement dynamics. Based on the neuronal connectivity and electrophysiology of the olivo-cerebellar network we have developed a general-purpose control approach, which we refer to as a universal control system (UCS), capable of dealing with a large number of actuator parameters in real time. In this UCS, the imposed goal and the resultant feedback from the actuators specify system properties. The goal is realized through implementing an architecture that can regulate a large number of parameters simultaneously by providing stimuli-modulated spatio-temporal cluster dynamics.

Spiking patterns emerging from wave instabilities in a one-dimensional neural lattice.

The dynamics of a one-dimensional lattice (chain) of electrically coupled neurons modeled by the FitzHugh-Nagumo excitable system with modified nonlinearity is investigated. We have found that for certain conditions the lattice exhibits a countable set of pulselike wave solutions. The analysis of homoclinic and heteroclinic bifurcations is given. Corresponding bifurcation sets have the shapes of spirals twisting to the same center. The appearance of chaotic spiking patterns emerging from wave instabilities is discussed.

Selective communication and information processing by excitable systems.

The phenomena of selective response of an excitable system to external pulse stimulation relating to interneuron communication and information processing problems are discussed. Subthreshold dynamics of the FitzHugh-Nagumo-like excitable system modeling of a neuron with the synaptic input is investigated. It is shown that the system response on various incoming information messages can be described by one- and two-dimensional linear and nonlinear point maps. Nonlinear integrating and resonant properties of the system are analyzed.

Theoretical and experimental study of two discrete coupled Nagumo chains.

We analyze front wave (kink and antikink) propagation and pattern formation in a system composed of two coupled discrete Nagumo chains using analytical and numerical methods. In the case of homogeneous interaction among the chains, we show the possibility of the effective control on wave propagation. In addition, physical experiments on electrical chains confirm all theoretical behaviors.

Synchronization, re-entry, and failure of spiral waves in a two-layer discrete excitable system.

A three-dimensional structure composed of two coupled discrete excitable lattices is considered. Each lattice (layer) is a discrete excitable subsystem and using a local model of excitation transfer and failure we have estimated the sufficient conditions for it to exhibit spiral waves. Then we show how interlayer synchronization of all motions is possible. Various effects of spiral wave synchronization, re-entry and failure are also investigated.

The emergence of form by replication.

It is shown with a simple mathematical model that if a system exhibits a given form (a spatial structure) and is put in contact with another system of the same type but in a state of spatial disorder, then under certain conditions their mutual interaction as they evolve in time allows replication of form in the disordered system with a controllable degree of faithfulness.