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The damage to dendrites and spines would result in decreased neuronal activity. Therefore, the decreased LFP response following 0. Hence, our results may provide some indication for the use of tPCS in animal and human studies. The calcium response can be reliably detected in neurogliopil region, which contains many fine processes of the surrounding neurons and astrocytes.

In these places, astrocyte processes closely interact with neuronal synapses and modulate synaptic transmission and plasticity Araque et al. The brain function arises from the coordinated activity of neuron-glia networks Perea et al. Many studies reported that tPCS can modulate brain oscillatory activity and enhance functional connectivity Guleyupoglu et al.

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Due to the different recording methods, the calcium activity observed in our experiments cannot be directly compared with their results, which were mainly based on EEG recording. In short, we cannot rule out the possibility that the neurogliopil response may also play a role in a-tPCS-induced plasticity. The findings in this study should be considered in the context of several limitations. Firstly, we acknowledge that the IP 3 R2 inositol trisphosphate receptor type 2 knockout mice should be used to prove the involvement of astrocytic GPCR G-protein-coupled receptor activation Monai et al.

However, the pharmacological experiments using prazosin demonstrated the involvement of noradrenergic activation of A1ARs, which transduce the G q signaling cascade for production of IP 3. Some related studies have demonstrated that synaptic plasticity in layer 4 disappears as the animal matures after the critical period Daw et al. In this study, the depth of imaging is limited to the superficial layers layers 1—3 of the cortex. Therefore, the findings in the current study are only pertinent to the superficial layers of the cortex.

Thirdly, the anodal current stimulation generally enhances cortical excitability. However, the a-tPCS involves the injection of much monopolar current. Compared to a-tPCS, transcranial alternating current stimulation tACS is a balanced current consisting of bipolar alternating current with equal electric charge.

No irreversible electrochemical products are known to accumulate at the electrode Antal et al.

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When considering translational validity in humans, a balanced tACS protocol might be an alternative. In short, the a-tPCS parameters in our study should be adjusted carefully when directly translated into the clinical conditions.

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In summary, the calcium response in mouse somatosensory cortex evoked by long-duration a-tPCS was reported for the first time. The decrease of cortical excitability induced by high-intensity a-tPCS may be caused by excessive calcium activity in neurons. These findings would contribute to the understanding of mechanisms underlying a-tPCS-induced cortical plasticity, and also suggest that the appropriate current intensity should be used in the application of a-tPCS. ZM and HS designed the experiments. ZM and HS wrote the manuscript. All authors reviewed the manuscript.

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. Alon, G. Safety and immediate effect of noninvasive transcranial pulsed current stimulation on gait and balance in Parkinson disease. Neural Repair 26, — Antal, A.

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Low intensity transcranial electric stimulation: safety, ethical, legal regulatory and application guidelines. Araque, A. Gliotransmitters travel in time and space. Neuron 81, — Arundine, M. Molecular mechanisms of calcium-dependent neurodegeneration in excitotoxicity. Cell Calcium 34, — Bastani, A. Differential modulation of corticospinal excitability by different current densities of anodal transcranial direct current stimulation.

Plos One 8:e Batsikadze, G. Partially non-linear stimulation intensity-dependent effects of direct current stimulation on motor cortex excitability in humans. Cambiaghi, M. Brain transcranial direct current stimulation modulates motor excitability in mice. Chung, W. Astrocytes control synapse formation, function, and elimination. Cold Spring Harbor Perspect.

Daw, N. Critical period for monocular deprivation in the cat visual cortex. Di Castro, M.

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Dissanayaka, T. Does transcranial electrical stimulation enhance corticospinal excitability of the motor cortex in healthy individuals? A systematic review and meta-analysis. Fitzgerald, P.

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Transcranial pulsed current stimulation: a new way forward? Fox, K. A critical period for experience-dependent synaptic plasticity in rat barrel cortex. Fregni, F. Regulatory considerations for the clinical and research use of transcranial direct current stimulation tDCS : review and recommendations from an expert panel. Fritsch, B. Direct current stimulation promotes BDNF-dependent synaptic plasticity: potential implications for motor learning.

Neuron 66, — Garaschuk, O. Targeted bulk-loading of fluorescent indicators for two-photon brain imaging in vivo. Guerra-Gomes, S. Functional roles of astrocyte calcium elevations: from synapses to behavior. Guleyupoglu, B.

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Classification of methods in transcranial electrical stimulation tES and evolving strategy from historical approaches to contemporary innovations. Methods , — Haydon, P. How do astrocytes participate in neural plasticity? Cold Spring Harb. Heller, J. Morphological plasticity of astroglia: understanding synaptic microenvironment. Glia 63, — Histed, M.

Direct activation of sparse, distributed populations of cortical neurons by electrical microstimulation. Neuron 63, — Jaberzadeh, S. Anodal transcranial pulsed current stimulation: a novel technique to enhance corticospinal excitability.

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Anodal transcranial pulsed current stimulation: the effects of pulse duration on corticospinal excitability. Plos One e Kabakov, A. Contribution of axonal orientation to pathway-dependent modulation of excitatory transmission by direct current stimulation in isolated rat hippocampus. Lalo, U. Exocytosis of gliotransmitters from cortical astrocytes: implications for synaptic plasticity and aging. Michelson, N.