activity of ZnO nanoparticles has been tested agains t some Gram-po sitive bacter ia such as Bacillus subtilis and Staphylococcus aureus, which have presented sensitivity to these nanoparticles (Adams et al., 2006; Gordon et al., 2011; Reddy et al., 2007). Also, ZnO has shown antimicrobial activity against Gram-negative bacteria such as Pseudomonas aeruginosa,Campylobacter jejuni and Escherichia coli (Brayner et al., 2006; Ohira et al., 2008; Premanathan et al., 2011; Sawai, 2003; Xie et al., 2011).E. coli has shown higher susceptibility to ZnO nanoparticles compared to S. aureus (Applerot et al., 2009a; Yamamoto, 2001). According to Applerot et al. (2009a), the higher resistance of S. aureus to ZnO nanoparticles can be explained by the differences between these two bacteriarelated to the intracellular antioxidant content such as carotenoid pigments in the interior ofS. aureus, which promote a greater oxidant resistance as well as the presence of potentdetoxification agents such as antioxidant enzymes, particularly catalase.However, the increased sensitivity of S. aureus to ZnO nanoparticles has also been reported (Adams et al., 2006; Premanathan et al., 2011; Reddy et al., 2007; Sawai, 2003). Accordingly, Sawai (2003) has suggested a strong affinity between ZnO nanoparticles and the bacteria cells of S. aureus as the cause of higher activity against this microorganism.There are two conditions for synergy between ZnO and S. aureus: the affinity of ZnO to the membrane of S. aureus and this microorganism's sensitivity to stress caused by H2O2 (Ohira et al. (2008). Previous work has indicated that ZnO nanoparticles generate H2O2.According to Russell (2003), Gram-negative bacteria have shown less sensitivity to reactive oxygen species (ROS) when compared with Gram-positive bacteria. One of the main reasons for this higher resistance is the structural differences in the bacterial membrane (Fig. 4).
Gram-positive bacteria have a membrane, which surrounds the cell, and a cell wall primarily made up of peptidoglycan layer as well as teichoic and lipoteichoic acids.
The cell wall of Gram-negative bacteria is more complex due to the presence of an outer membrane, which is composed mainly of lipopolysaccharide (LPS), in addition to a
thin peptidoglycan layer (Epand & Epand, 2009; Jiang et al., 2004). Thus, the outer membrane of Gram-negative bacteria acts as a permeability barrier, so that the absorption of ROS into the cell is reduced (Russell, 2003).
Another explanation for the increased resistance to ZnO activity observed in E. coli compared with S. aureus is due to differences in the polarity of the cell membrane, since the membrane ofS. aureus has less negative charge than E. coli (Sonohara et al., 1995). According to Gordon et al. (2011), this would allow a greater penetration level of negatively charged free radicals such as hydroxyl radicals, superoxide and peroxide ions causing damage and cell death in S. aureus at concentrations below that required to cause the same effect in E. coli.
Although several mechanisms have been proposed to explain the differences in the antimicrobial activity of ZnO
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