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However, for such experiments to proceed and succeed, more information is needed regarding signal transmission across the gap junctions associated with astrocytes. The AN model also shows that astrocytes have a key role in the dynamic synchronization of neurons.

The SICs and neurotransmitter modulation induced by astrocytic release of glutamate causes coordinated neural activity in remote neurons. These should be out of phase due to the different initiation times of the input spike trains. The latter arises from modulation of neurotransmitter in the cleft of all synapses associated with the astrocyte and consequently the IP 3 level in the astrocyte. Moreover, it has recently been suggested that oscillations within microdomains of astrocytic distal processes differ from oscillations at the soma []. Despite this simplistic assumption our results show that when phase locking cannot be achieved, oscillations within the distal processes and the soma are indeed different.

Our results also suggest that even though phase locking can occur it is not vital for coordination. However, when both input stimulus frequencies were below 12 Hz and again in antiphase, coordination was still possible. Consequently, no further global release of glutamate from the astrocyte was possible and the SICs died off. Although propagation delays were not included in the present version of the AN model it would be interesting to investigate intercellular delays and the propagation time for waves across astrocyte networks.

For example, it is likely that dynamic coordination across neuron clusters, mediated via astrocyte networks, is important for the brain rhythms underlying cognitive function. Moreover, the phase locking characteristics of our model may also provide a mechanism for dynamically changing the coordination between neuron ensembles.

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Since phase locking was found to occur between microdomain oscillations with approximately the same frequency of oscillation, it is not inconceivable that stimuli of similar frequency may cause a unique pattern of oscillation which changes as the stimuli frequencies change.

Furthermore, this threshold may also have an impact on the pattern of coordination between neurons. This condition may be avoided, however, if the threshold is raised. More biological experimentation is required to establish the key parameters governing the threshold level.

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This finding supports the recent suggestion that these infra-slow oscillations are of astrocytic origin [] , []. Since the pattern of coordination observed here is based primarily on independent synaptic stimuli, one role of phase synchronization may be information encoding and communication [27]. Astrocytes, like neurons, also form interconnected networks where the communication between astrocytes is via gap junctions.

It is worth noting that the various model systems encompassed by the AN model were chosen to reduce computational overhead while still remaining biophysically meaningful. Thus, the AN model can form an important building block to explore global communication across large remote clusters of neurons via astrocyte ensembles. Extension of the AN model to include intercellular signaling similar to those described in [] — [] may aid in the understanding of how phase locking of field potentials encodes functionally relevant information via AN networks.

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The present modeling results strengthen the hypothesis that astrocyte networks provide much more than structural support to neural networks. Indeed astrocytes are viewed as regulators of neural circuitry through coordination of transmission at synaptic junctions. Moreover, it is also believed that retrograde messengers induced in the postsynaptic neuron can be fed back either directly or via an astrocyte to receptors on the presynaptic neuron []. This feedback signal modulates the probability of neurotransmitter release and therefore may provide new insight into self repairing synapses Extension of the present AN model may thus be used to investigate such repair mechanisms [].

Moreover, the AN model could be extended to provide a flexible research tool to allow neuroscientists to explore the role of astrocytes in a number of neurological disorders. IP 3 generation within the astrocyte cytoplasm. A The evolution of IP 3 within the cytoplasm of the astrocyte as a result of a range of Poisson generated spike trains stimulating the tripartite synapse. Note that IP 3 builds much faster than it decays which can be seen after 10 s when the input ceases. B Same experiment as A except the Poisson distributed spike train is maintained. Note how the level of generated IP 3 is limited to a steady state value and is dependent on the stimulus frequency.

A AM mode with input stimulus frequency set at 3 Hz. B FM mode with input stimulus set at 7 Hz. A AM mode with the input stimulus frequency set at 18 Hz. B FM mode with the input stimulus frequency set at 36 Hz.

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  • As a result IP 3 reaches a steady state at which no further crossing of the threshold is possible. The authors are grateful to Dr. Suhita Nadkarni for helpful and insightful comments that improved the paper considerably. Performed the experiments: JJW. Browse Subject Areas? Click through the PLOS taxonomy to find articles in your field. Abstract In recent years research suggests that astrocyte networks, in addition to nutrient and waste processing functions, regulate both structural and synaptic plasticity.

    Introduction For many years, astrocytes, a subgroup of glial cells found in the brain, have been thought to support neurons by providing them with vital elements needed for their survival [1] — [3]. Materials and Methods A feature of the present modeling approach is its constructive nature: it combines and constructs multiple detailed models in order to reveal the regulatory dynamics of astrocytes at a network level. Astrocyte — Neuron Interactions Both the gatekeeper [30] and Nadkarni and Jung [31] , [32] models describe the interaction of astrocytes and neurons via the tripartite synapse.

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    Synapse model and astrocyte feedback Synaptic information transfer is considered to be probabilistic: on arrival of an AP a vesicle is either released or it is not [41]. Neuron Model Although many neuron models exist [53] such as the Hodgkin-Huxley model [54] simplified counterparts such as the FitzHugh-Nagumo [55] , [56] and Morris-Lecar models [57] are often preferred by electrophysiologists [53].

    Plasticity Model Since Donald Hebb first suggested that a neuron must fire shortly before or at the same time as a neuron to which it is connected in order to strengthen the connection between them [64] , [65] , there have been many mathematical models that explore synaptic plasticity see [66] for review. Figure 2. AN Model Block Diagram showing interactions between an astrocyte and neuron cell.

    Results In all the results presented here, Matlab a platform Windows version by Mathworks was used to realize the AN model and the Euler method of integration was used for simulation. Establishing the Valid Range of Input Frequencies In this simulation, presynaptic neuron A stimulates a tripartite synapse with a sustained Poisson spike train for s see Figure 3. Figure 3.

    Network consists of presynaptic neuron A, postsynaptic neuron B and an interconnecting tripartite synapse. Figure 4. Spatially distributed learning signals Here we show how a learning signal, distributed spatially via an astrocyte, can promote STDP-based plasticity at remote synaptic sites.

    Figure 8. Synaptic Plasticity in the AN Model 2. Dynamic Coordination Neural oscillations across a broad range of frequency bands are ubiquitous throughout the nervous system and give rise to a wide variety of dynamic coordination effects including synchrony, learning, precisely timed phase-shifts among oscillating neural ensembles, concatenation of different rhythms, and so forth [92] — [96]. Figure Coordinated firing activity of N1 and N2 for the first 15 s of AN model simulation. Discussion The AN model presented in this paper captures the bidirectional coupling between astrocytes and neurons and in so doing demonstrates that positive and negative feedback to extrasynaptic NMDARs and presynaptic mGluRs significantly contributes to the regulatory capability of astrocytes.

    Dynamic coordination The AN model also shows that astrocytes have a key role in the dynamic synchronization of neurons. Extension of the AN model Astrocytes, like neurons, also form interconnected networks where the communication between astrocytes is via gap junctions.

    Supporting Information. Figure S1.

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    Figure S2. Figure S3. Acknowledgments The authors are grateful to Dr. References 1. View Article Google Scholar 2. Nat Neurosci 1: — View Article Google Scholar 3. Kurosinski P, Gotz J Glial cells under physiologic and pathologic conditions. Arch Neurol — View Article Google Scholar 4. J Neurosci — View Article Google Scholar 5. View Article Google Scholar 6. Neuron 8: — View Article Google Scholar 7. Zur Nieden R, Deitmer JW The role of metabotropic glutamate receptors for the generation of calcium oscillations in rat hippocampal astrocytes in situ.

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