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      Are Nanoparticles a Threat to Mycorrhizal and Rhizobial Symbioses? A Critical Review

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          Abstract

          Soil microorganisms can be exposed to, and affected by, nanoparticles (NPs) that are either purposely released into the environment (e.g., nanoagrochemicals and NP-containing amendments) or reach soil as nanomaterial contaminants. It is crucial to evaluate the potential impact of NPs on key plant-microbe symbioses such as mycorrhizas and rhizobia, which are vital for health, functioning and sustainability of both natural and agricultural ecosystems. Our critical review of the literature indicates that NPs may have neutral, negative, or positive effects on development of mycorrhizal and rhizobial symbioses. The net effect of NPs on mycorrhizal development is driven by various factors including NPs type, speciation, size, concentration, fungal species, and soil physicochemical properties. As expected for potentially toxic substances, NPs concentration was found to be the most critical factor determining the toxicity of NPs against mycorrhizas, as even less toxic NPs such as ZnO NPs can be inhibitory at high concentrations, and highly toxic NPs such as Ag NPs can be stimulatory at low concentrations. Likewise, rhizobia show differential responses to NPs depending on the NPs concentration and the properties of NPs, rhizobia, and growth substrate, however, most rhizobial studies have been conducted in soil-less media, and the documented effects cannot be simply interpreted within soil systems in which complex interactions occur. Overall, most studies indicating adverse effects of NPs on mycorrhizas and rhizobia have been performed using either unrealistically high NP concentrations that are unlikely to occur in soil, or simple soil-less media (e.g., hydroponic cultures) that provide limited information about the processes occurring in the real environment/agrosystems. To safeguard these ecologically paramount associations, along with other ecotoxicological considerations, large-scale application of NPs in farming systems should be preceded by long-term field trials and requires an appropriate application rate and comprehensive (preferably case-specific) assessment of the context parameters i.e., the properties of NPs, microbial symbionts, and soil. Directions and priorities for future research are proposed based on the gaps and experimental restrictions identified.

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          Most cited references108

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          Towards a definition of inorganic nanoparticles from an environmental, health and safety perspective.

          The regulation of engineered nanoparticles requires a widely agreed definition of such particles. Nanoparticles are routinely defined as particles with sizes between about 1 and 100 nm that show properties that are not found in bulk samples of the same material. Here we argue that evidence for novel size-dependent properties alone, rather than particle size, should be the primary criterion in any definition of nanoparticles when making decisions about their regulation for environmental, health and safety reasons. We review the size-dependent properties of a variety of inorganic nanoparticles and find that particles larger than about 30 nm do not in general show properties that would require regulatory scrutiny beyond that required for their bulk counterparts.
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            Coordinating nodule morphogenesis with rhizobial infection in legumes.

            The formation of nitrogen-fixing nodules on legumes requires an integration of infection by rhizobia at the root epidermis and the initiation of cell division in the cortex, several cell layers away from the sites of infection. Several recent developments have added to our understanding of the signaling events in the epidermis associated with the perception of rhizobial nodulation factors and the role of plant hormones in the activation of cell division leading to nodule morphogenesis. This review focuses on the tissue-specific nature of the developmental processes associated with nodulation and the mechanisms by which these processes are coordinated during the formation of a nodule.
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              Oxidative stress-mediated antibacterial activity of graphene oxide and reduced graphene oxide in Pseudomonas aeruginosa

              Background Graphene holds great promise for potential use in next-generation electronic and photonic devices due to its unique high carrier mobility, good optical transparency, large surface area, and biocompatibility. The aim of this study was to investigate the antibacterial effects of graphene oxide (GO) and reduced graphene oxide (rGO) in Pseudomonas aeruginosa. In this work, we used a novel reducing agent, betamercaptoethanol (BME), for synthesis of graphene to avoid the use of toxic materials. To uncover the impacts of GO and rGO on human health, the antibacterial activity of two types of graphene-based material toward a bacterial model P. aeruginosa was studied and compared. Methods The synthesized GO and rGO was characterized by ultraviolet-visible absorption spectroscopy, particle-size analyzer, X-ray diffraction, scanning electron microscopy and Raman spectroscopy. Further, to explain the antimicrobial activity of graphene oxide and reduced graphene oxide, we employed various assays, such as cell growth, cell viability, reactive oxygen species generation, and DNA fragmentation. Results Ultraviolet-visible spectra of the samples confirmed the transition of GO into graphene. Dynamic light-scattering analyses showed the average size among the two types of graphene materials. X-ray diffraction data validated the structure of graphene sheets, and high-resolution scanning electron microscopy was employed to investigate the morphologies of prepared graphene. Raman spectroscopy data indicated the removal of oxygen-containing functional groups from the surface of GO and the formation of graphene. The exposure of cells to GO and rGO induced the production of superoxide radical anion and loss of cell viability. Results suggest that the antibacterial activities are contributed to by loss of cell viability, induced oxidative stress, and DNA fragmentation. Conclusion The antibacterial activities of GO and rGO against P. aeruginosa were compared. The loss of P. aeruginosa viability increased in a dose- and time-dependent manner. Exposure to GO and rGO induced significant production of superoxide radical anion compared to control. GO and rGO showed dose-dependent antibacterial activity against P. aeruginosa cells through the generation of reactive oxygen species, leading to cell death, which was further confirmed through resulting nuclear fragmentation. The data presented here are novel in that they prove that GO and rGO are effective bactericidal agents against P. aeruginosa, which would be used as a future antibacterial agent.
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                Author and article information

                Contributors
                Journal
                Front Microbiol
                Front Microbiol
                Front. Microbiol.
                Frontiers in Microbiology
                Frontiers Media S.A.
                1664-302X
                24 July 2019
                2019
                : 10
                : 1660
                Affiliations
                [1] 1Key Laboratory of Plant Nutrition and Agri-environment in Northwest China, Ministry of Agriculture, College of Natural Resources and Environment, Northwest A&F University , Yangling, China
                [2] 2School of Environment, The University of Auckland , Auckland, New Zealand
                [3] 3School of Agriculture and Environment, The University of Western Australia , Crawley, WA, Australia
                Author notes

                Edited by: Saskia Bindschedler, Université de Neuchâtel, Switzerland

                Reviewed by: Alison Bennett, The Ohio State University, United States; Muhammad Saleem, Alabama State University, United States

                *Correspondence: Hui Tian, tianh@ 123456nwsuaf.edu.cn

                This article was submitted to Terrestrial Microbiology, a section of the journal Frontiers in Microbiology

                Article
                10.3389/fmicb.2019.01660
                6668500
                31396182
                69fd32b8-d814-4283-9434-bdbeb78e45ef
                Copyright © 2019 Tian, Kah and Kariman.

                This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

                History
                : 27 February 2019
                : 04 July 2019
                Page count
                Figures: 3, Tables: 3, Equations: 0, References: 122, Pages: 15, Words: 0
                Categories
                Microbiology
                Review

                Microbiology & Virology
                nanoparticles,soil,mycorrhiza,rhizobia,colonization,nodule,toxicity
                Microbiology & Virology
                nanoparticles, soil, mycorrhiza, rhizobia, colonization, nodule, toxicity

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