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Theranostics 2020, Vol. 10, Issue 20 concentration-dependent manner, as well as the increased number of G0/G1 phase cells, which may be prone to apoptosis [244, 258, 261]. AgNPs can not only induce apoptosis through ROS-mediated DNA damage, but also destroy DNA structure directly via Ag0 and Ag+ released by AgNPs [157]. The DNA double helix structure is composed of four bases of adenine, guanine, cytosine and thymine by strictly complementary base pairing. Base pairs are bounded by hydrogen bonds to form a unit of DNA double helix. The destruction of hydrogen bonds decreases the stability of DNA structure. Tsuneo Ishida [157] analyzed the activities of AgNPs in the nucleus. Silver could form a complex containing silver within DNA. Ag+ caused DNA damage by replacing the hydrogen bonds in the G≡C and A=T base pairs. The Ag atom was twofold coordinated by two N atoms to form N-Ag+-N complex in G≡C base pair, and other complex structures appearing in the base pair were O-Ag+-N (G≡C base pair), N-Ag+-O (both G≡C and A=T base pairs). DNA damage caused by these complexes might be a factor in triggering cancer cell apoptosis. Generally speaking, AgNPs can exert anti-cancer effects through multiple pathways. Bandyopadhyay et al. [262] confirmed that AgNPs could exhibit antitumor properties through multiple channels, including triggering cell morphological changes, ROS generation, and nuclear fragmentation, while exhibited minimum toxicity in normal peripheral blood lymphocytes. The considerable anticancer activity and histocompatibility might relate to the types of reducing agent and stabilizer. Inactivate Proteins and Regulate Signaling Pathways In the development and progression of tumors, many signaling pathways are involved in controlling cell growth and proliferation, apoptosis and viability, and can participate in more complex signaling networks that contribute to tumor progression, such as tumor microenvironment (TME), angiogenesis and inflammation [263]. Some proteases and cytokines are also involved in these regulations, such as vascular endothelial growth factor (VEGF), matrix metalloproteinase (MMPs) and fibroblast growth factor 2 (FGF-2), etc. AgNPs have been confirmed to inhibit tumor proliferation, invasion and angiogenesis by regulating the associated signaling pathways or inactivating proteins. For example, AgNPs can regulate the HIF signaling pathway [161]. In general, rapid proliferation of tumor cells and irregular vasculature cause hypoxic TME [264-266]. HIF-1 level is up-regulated in hypoxic TME, followed by activation of target genes that in response to hypoxia. These genes contribute to cellular activities, such as 9013 cell proliferation, angiogenesis, and eventually lead to failure of cancer treatment [161]. Therefore, HIF-1 is a potential target for cancer treatment. It has been demonstrated that hypoxia can weaken HIF-1α- mediated autophagy [247]. Tieshan Yang et al. found that AgNPs could disrupt the HIF signaling pathway by attenuating HIF-1 protein accumulation and downstream target genes expression [161]. AgNPs can also inhibit the progression of tumors by inhibiting MMPs activity. MMPs are known as protein family and classified into different evolutionary groups according to their primary sequences [267]. MMPs play a dominant role in tumor progressions, such as tumor cell proliferation, invasiveness and distant metastasis, evasion of immune surveillance, and angiogenesis [267, 268]. Therefore, MMPs are considered as potential targets for cancer therapy [31]. In order to obtain antitumor drugs with targeting capabilities, some teams have attempted to develop inhibitors against members of MMPs. Other signaling pathways and proteases involved in tumor progression have also been highlighted. Melissa M Kemp et al. [245] found that AgNPs could effectively inhibit FGF-2-induced angiogenesis. Their results suggested that AgNPs may have great potential for inhibiting pathological angiogenesis in cancer. Eom et al. [27] indicated that AgNPs induced cytotoxicity, including DNA damage, cell cycle arrest and apoptosis, by activating the p38 MAPK signaling cascades. These studies may inspire the development of anticancer agents containing AgNPs. In view of the complex signaling pathways and various proteins involved in the regulation of tumor development and progression, anticancer mechanisms of AgNPs by regulating intracellular signaling pathways and inactivating proteins still need to be further explored. Inhibit Migration and Angiogenesis Numerous studies have confirmed that AgNPs can inhibit migration and invasion of tumor cells in concentration- and dose-dependent manners [23, 30, 32, 269]. Migration and invasion are important hallmarks of cancer progression and deterioration [270]. Although it has been observed that AgNPs can inhibit tumor invasion [269], the specific mechanism is still unclear. It is hypothesized that AgNPs may decrease the protein expression of cytokines and growth factors within cancer cells, or reduce the enzymatic activity of MMPs. VEGF is an important signaling protein involved in vasculogenesis and angiogenesis, which plays a crucial role in tumor growth and metastasis [32]. Various studies support that AgNPs can deprive cancer cells of both nutrients http://www.thno.orgPDF Image | Silver nanoparticles Synthesis medical applications safety
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