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and increased glucose uptake, irrespective of oxygen availability (aerobic glycolysis or Warburg effect) [52], we believe that they are able to detect the sugars present in the EPS and to pick it with greater efficiency compared to normal cells. Recently, carbohydrates have been considered as biomimetic functional molecules on the surface of nanoparticles [53] also because they trigger cellular uptake via specific receptors or endocytosis. TEM analysis showed that AgNPs-EPSaer uptake occurs through endocytosis and, as suggested by colony formation assay, the effects are evident after 1h of treatment. Moreover, AgNPs-EPSaer treatment inhibited migration and invasion capability of tumoral cells, considered as a cancer hallmark, suggesting a potential anticancer effect. Findings from several studies reported that both the AgNPs and Ag+ released by AgNPs are involved in the mechanism of cytotoxicity in different ways: infact, AgNPs probably provide a perfect surface outside the mitochondria for the univalent reduction of oxygen to superoxide from electron through the electron transport chain. On the other hand, Ag+ binds to proteins and DNA, interfering with their functions [54]. Moreover, ROS generation and oxidative stress occur as an early event leading to NP-induced toxicity [38]. Our results point in the same direction suggesting that the cellular response to AgNPs-EPSaer exposure is the sum of events triggered by a direct effect of the cellular uptake of AgNPs, as verified by TEM analysis, induction of intracellular ROS and an indirect effect of free Ag+ ions released mainly in mitochondria and secondarily in the nuclei, where they interact with DNA. To the best of our knowledge this is the first study reporting quali- quantitative evidences about the effects of siver ions released by nanoparticles in different cell compartments. It is generally accepted that ROS induce autophagy and/or apoptosis. Moreover, it has been documented that autophagy may act as enabler of apoptosis, contributing in certain morphological and cellular events that take place in apoptotic cell death [55]. Autophagic cell death has morphologic and biochemical features distinguishing it from both apoptosis and necrosis. Autophagy consists of several sequential steps starting with the autophagosome formation and then progresses to autophagolysosomes through the fusion of acidic lysosomes with autophagosomes [56–58]. Several proteins can be used as markers to study autophagic flux: in particular, during autophagy, ATG proteins (eg ATG5 and ATG7) and Beclin -1, have an essential role in the autophagosome formation [59]. Later, autophagosomes (LC3-containing vacuoles) fuse with lysosomes to cause the degradation of their contents. The protein p62/SQSTM1 binds directly to LC3 and is itself degraded [60, 61]. Moreover, inhibition of HSP90 plays an important role in autophagy inducing degradation of Hsp90 client proteins, such as AKT [62, 63]. PI3K/AKT/mTOR axis downregulation lead to the activation of autophagy and the inhibition of cell proliferation, also [63]. We found that SKBR3 treated with AgNPs-EPSaer underwent morphologic and biochemical changes consistent with the induction of autophagy and secondarily of apoptosis, as suggested by the upregulation of lysosomes, autophagolysosomes and Figure 10: Determination of total silver by ICP in cell fractions. Data were normalized for protein concentration. ND means not detected. www.impactjournals.com/oncotarget 9696 OncotargetPDF Image | Anticancer activity of biogenerated silver nanoparticles
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