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Controlling stem cell fate using cold atmospheric plasma Fei Tan1,2,3* , Yin Fang2, Liwei Zhu1 and Mohamed Al-Rubeai4

Abstract The stem cell is the foundation of regenerative medicine and tissue engineering. Regulating specific stem cell fate, such as cell attachment, proliferation, differentiation, and even death, undergoes continuous development. Cold atmospheric plasma (CAP), the core technology of plasma medicine, is attracting tremendous attention due to its ability and versatility to manipulate various types of cells, including stem cells. Specifically, the direct and indirect applications of CAP in controlling cell fate are best exemplified by upfront irradiation of the stem cells and modification of the stem cell niche, respectively. This review will describe the recent advances in various CAP strategies, both direct and indirect, and their influence on the fate of healthy and cancer stem cells. Particular emphasis will be placed on the mechanism of connecting the physical and chemical cues carried by the plasma and biological changes presented by the cells, especially at the transcriptomic level. The ultimate goal is to exploit CAP’s potential in regenerative medicine. Keywords: Atmospheric plasma, Stem cell, Cell attachment, Cell proliferation, Cell differentiation, Plasma medicine, Regenerative medicine, Cell death, Tissue engineering, Cold plasma, Non-thermal plasma, Stem cell niche, Surface modification, Scaffold, Extracellular matrix, Plasma-activated medium

Cold atmospheric plasma In physics, plasma is the so-called fourth state of matter consisting of roughly equal numbers of positively and negatively charged particles. It could be produced typically at very high temperature or at low pressures and acquired either naturally or artificially. Cold atmospheric plasma (CAP) is a partially ionized gas generated at atmospheric pressure and operates under room temperature. This term is sometimes interchangeable with low-temperature plasma (LTP), non-thermal atmospheric plasma (NTAP), and non-equilibrium atmospheric plasma (NAP). Various technologies have been used to generate atmospheric plasma (Fig. 1), including, but are not limited to, pulsed atmospheric arc (PAA) technology (Fig. 1a) and * Correspondence: [email protected] 1 Department of ORL-HNS, Affiliated East Hospital of Tongji University, Shanghai, China 2 School of Medicine and Institute for Advanced Study, Tongji University, Shanghai, China Full list of author information is available at the end of the article

piezoelectric direct discharge (PDD) technology (Fig. 1b), each with distinct advantages and disadvantages. A detailed comparison between these two technologies and their medical applicability can be found in Table 1. The key components of CAP include reactive oxygen and nitrogen species (RONS), ions and electrons, and UV photons [1]. The composition and concentration of these components can be tuned and programmed for various applications, especially biomedical ones. In order for CAP to directly influence living