Pulse coupled oscillators and the phase resetting curve (original) (raw)
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Phase resetting and phase locking in hybrid circuits of one model and one biological neuron
Biophysical journal, 2004
To determine why elements of central pattern generators phase lock in a particular pattern under some conditions but not others, we tested a theoretical pattern prediction method. The method is based on the tabulated open loop pulsatile interactions of bursting neurons on a cycle-by-cycle basis and was tested in closed loop hybrid circuits composed of one bursting biological neuron and one bursting model neuron coupled using the dynamic clamp. A total of 164 hybrid networks were formed by varying the synaptic conductances. The prediction of 1:1 phase locking agreed qualitatively with the experimental observations, except in three hybrid circuits in which 1:1 locking was predicted but not observed. Correct predictions sometimes required consideration of the second order phase resetting, which measures the change in the timing of the second burst after the perturbation. The method was robust to offsets between the initiation of bursting in the presynaptic neuron and the activation of the synaptic coupling with the postsynaptic neuron. The quantitative accuracy of the predictions fell within the variability (10%) in the experimentally observed intrinsic period and phase resetting curve (PRC), despite changes in the burst duration of the neurons between open and closed loop conditions.
Biophysical journal, 2009
Existence and stability criteria for harmonic locking modes were derived for two reciprocally pulse coupled oscillators based on their first and second order phase resetting curves. Our theoretical methods are general in the sense that no assumptions about the strength of coupling, type of synaptic coupling, and model are made. These methods were then tested using two reciprocally inhibitory Wang and Buzsáki model neurons. The existence of bands of 2:1, 3:1, 4:1, and 5:1 phase locking in the relative frequency parameter space was predicted correctly, as was the phase of the slow neuron's spike within the cycle of the fast neuron in which it occurred. For weak coupling the bands are very narrow, but strong coupling broadens the bands. The predictions of the pulse coupled method agreed with weak coupling methods in the weak coupling regime, but extended predictability into the strong coupling regime. We show that our prediction method generalizes to pairs of neural oscillators coupled through excitatory synapses, and to networks of multiple oscillatory neurons. The main limitation of the method is the central assumption that the effect of each input dies out before the next input is received.
Journal of Computational Neuroscience, 2011
Gamma oscillations can synchronize with near zero phase lag over multiple cortical regions and between hemispheres, and between two distal sites in hippocampal slices. How synchronization can take place over long distances in a stable manner is considered an open question. The phase resetting curve (PRC) keeps track of how much an input advances or delays the next spike, depending upon where in the cycle it is received. We use PRCs under the assumption of pulsatile coupling to derive existence and stability criteria for 1:1 phase-locking that arises via bidirectional pulse coupling of two limit cycle oscillators with a conduction delay of any duration for any 1:1 firing pattern. The coupling can be strong as long as the effect of one input dissipates before the next input is received. We show the form that the generic synchronous and anti-phase solutions take in a system of two identical, identically pulse-coupled oscillators with identical delays. The stability criterion has a simple form that depends only on the slopes of the PRCs at the phases at which inputs are received and on the number of cycles required to complete the delayed feedback loop. The number of cycles required to complete the delayed feedback loop depends upon both the value of the delay and the firing pattern. We successfully tested the predictions of our methods on networks of model neurons. The criteria can easily be extended to include the effect of an input on the cycle after the one in which it is received.
Journal of Neuroscience, 2009
Networks of model neurons were constructed and their activity was predicted using an iterated map based solely on the phase-resetting curves (PRCs). The predictions were quite accurate provided that the resetting to simultaneous inputs was calculated using the sum of the simultaneously active conductances, obviating the need for weak coupling assumptions. Fully synchronous activity was observed only when the slope of the PRC at a phase of zero, corresponding to spike initiation, was positive. A novel stability criterion was developed and tested for all-to-all networks of identical, identically connected neurons. When the PRC generated using N Ϫ 1 simultaneously active inputs becomes too steep, the fully synchronous mode loses stability in a network of N model neurons. Therefore, the stability of synchrony can be lost by increasing the slope of this PRC either by increasing the network size or the strength of the individual synapses. Existence and stability criteria were also developed and tested for the splay mode in which neurons fire sequentially. Finally, N/M synchronous subclusters of M neurons were predicted using the intersection of parameters that supported both between-cluster splay and within-cluster synchrony. Surprisingly, the splay mode between clusters could enforce synchrony on subclusters that were incapable of synchronizing themselves. These results can be used to gain insights into the activity of networks of biological neurons whose PRCs can be measured.
Network of phase-locking oscillators and a possible model for neural synchronization
Communications in Nonlinear Science and Numerical Simulation, 2011
In order to model the synchronization of brain signals, a three-node fully-connected network is presented. The nodes are considered to be voltage control oscillator neurons (VCON) allowing to conjecture about how the whole process depends on synaptic gains, free-running frequencies and delays. The VCON, represented by phase-locked loops (PLL), are fully-connected and, as a consequence, an asymptotically stable synchronous state appears. Here, an expression for the synchronous state frequency is derived and the parameter dependence of its stability is discussed. Numerical simulations are performed providing conditions for the use of the derived formulae. Model differential equations are hard to be analytically treated, but some simplifying assumptions combined with simulations provide an alternative formulation for the long-term behavior of the fully-connected VCON network. Regarding this kind of network as models for brain frequency signal processing, with each PLL representing a neuron (VCON), conditions for their synchronization are proposed, considering the different bands of brain activity signals and relating them to synaptic gains, delays and free-running frequencies. For the delta waves, the synchronous state depends strongly on the delays. However, for alpha, beta and theta waves, the free-running individual frequencies determine the synchronous state.
Journal of Computational Neuroscience, 2008
Our goal is to understand how nearly synchronous modes arise in heterogenous networks of neurons. In heterogenous networks, instead of exact synchrony, nearly synchronous modes arise, which include both 1:1 and 2:2 phase-locked modes. Existence and stability criteria for 2:2 phase-locked modes in reciprocally coupled two neuron circuits were derived based on the open loop phase resetting curve (PRC) without the assumption of weak coupling. The PRC for each component neuron was generated using the change in synaptic conductance produced by a presynaptic action potential as the perturbation. Separate derivations were required for modes in which the firing order is preserved and for those in which it alternates. Networks composed of two model neurons coupled by reciprocal inhibition were examined to test the predictions. The parameter regimes in which both types of nearly synchronous modes are exhibited were accurately predicted both qualitatively and quantitatively provided that the synaptic time constant is short with respect to the period and that the effect of second order resetting is considered. In contrast, PRC methods based on weak coupling could not predict 2:2 modes and did not predict the 1:1 modes with the level of accuracy achieved by the strong coupling methods. The strong coupling prediction methods provide insight into what manipulations promote near-synchrony in a two neuron network and may also have predictive value for larger networks, which can also manifest changes in firing order. We also identify a novel route by which synchrony is lost in mildly heterogenous networks.
Lecture Notes in Computer Science, 2009
Fully-connected neural networks of non-linear half-center oscillators coupled both electrically and synaptically may exhibit a variety of modes of oscillation despite fixed topology and parameters. In suitable circumstances it is possible to switch between these modes in a controlled fashion. Previous work has investigated this phenomenon the simplest possible 2 cell network. In this paper we show that the 4 cell network, like the 2 cell, exhibits a variety of symmetrical and asymmetrical behaviours. In general, with increasing electrical coupling the number of possible behaviours is reduced until finally the only expressed behaviour becomes in-phase oscillation of all neurons. Our analysis enables us to predict general rules governing behaviour of more numerous networks, for instance types of co-existing activity patterns and a subspace of parameters where they emerge. 1 synchrony has two meanings: it signifies rhythmic activity expressed with the same frequency, all neurons therefore must be phase-locked with some phase (for example equal to 0-in-phase behaviour-or 0.5 of the cycle out-of-phase behaviour) or it signifies a phase locking equal 0 (i. e. in-phase behaviour). We will use here the term synchrony in its former, more general, meaning.
Phase Response Curves in Neuroscience, 2011
We use phase resetting curve (PRC) theory to analyze phase-locked patterns in pulse-coupled all to all network of N neurons that receive multiple inputs per cycle. The basic principles are that the phase must be updated each time an input is received, and simultaneous inputs do not sum linearly for strong coupling, but the conductances do. Therefore, the dependence of the resetting on conductance must be known. We analyze a splay mode in which no neurons fire simultaneously, global synchrony in which all neurons fire together, and clustering modes in which the firing breaks up into a small number of clusters. The key idea is to identify the appropriate perturbation in order to determine the stability of a given mode. For the splay mode, jitter is introduced into all firing times. For synchrony, a single neuron is perturbed from the rest, and for the two cluster mode, a single neuron is perturbed from one cluster. Global synchrony can be destabilized by increasing the network size or the strength of the individual synapses. At most, a small number of M clusters form because the M 1 locking points are more likely to sample destabilizing regions of the PRC as M increases. Between cluster interactions can enforce synchrony on subclusters that are incapable of synchronizing themselves. For the two cluster case, general results were obtained for clusters of any size. These results can be used to gain insights into the activity of networks of biological neurons whose PRCs can be measured.
PLoS ONE, 2014
We studied the dynamics of a large-scale model network comprised of oscillating electrically coupled neurons. Cells are modeled as relaxation oscillators with short duty cycle, so they can be considered either as models of pacemaker cells, spiking cells with fast regenerative and slow recovery variables or firing rate models of excitatory cells with synaptic depression or cellular adaptation. It was already shown that electrically coupled relaxation oscillators exhibit not only synchrony but also anti-phase behavior if electrical coupling is weak. We show that a much wider spectrum of spatiotemporal patterns of activity can emerge in a network of electrically coupled cells as a result of switching from synchrony, produced by short external signals of different spatial profiles. The variety of patterns increases with decreasing rate of neuronal firing (or duty cycle) and with decreasing strength of electrical coupling. We study also the effect of network topology -from all-to-all -to pure ring connectivity, where only the closest neighbors are coupled. We show that the ring topology promotes anti-phase behavior as compared to all-to-all coupling. It also gives rise to a hierarchical organization of activity: during each of the main phases of a given pattern cells fire in a particular sequence determined by the local connectivity. We have analyzed the behavior of the network using geometric phase plane methods and we give heuristic explanations of our findings. Our results show that complex spatiotemporal activity patterns can emerge due to the action of stochastic or sensory stimuli in neural networks without chemical synapses, where each cell is equally coupled to others via gap junctions. This suggests that in developing nervous systems where only electrical coupling is present such a mechanism can lead to the establishment of proto-networks generating premature multiphase oscillations whereas the subsequent emergence of chemical synapses would later stabilize generated patterns.
Phase-locked oscillations in a neuronal network model
Neurocomputing, 2002
We analyzed the oscillatory activities in a neuronal network model as the basis of synchrony of the activities in the brain. The model consists of two groups of neurons that are interconnected. One group is composed of an excitatory and an inhibitory neuron which are expressed by Hodgkin-Huxley equations. The network shows di erent phase-locked oscillations depending on the structure and intensity of interconnection between groups or coupling of neurons in the group, or the value of synaptic latency. The oscillations include various periodic solutions in which the two groups oscillate not only in in-phase or anti-phase but also in continuously changing phase di erence with the parameters of coupling and latency.