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      Volatile organic compounds emitted by Trichoderma species mediate plant growth

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          Abstract

          Background

          Many Trichoderma species are applied as biofungicides and biofertilizers to agricultural soils to enhance crop growth. These filamentous fungi have the ability to reduce plant diseases and promote plant growth and productivity through overlapping modes of action including induced systemic resistance, antibiosis, enhanced nutrient efficiency, and myco-parasitism. Trichoderma species are prolific producers of many small metabolites with antifungal, antibacterial, and anticancer properties. Volatile metabolites of Trichoderma also have the ability to induce resistance to plant pathogens leading to improved plant health. In this study, Arabidopsis plants were exposed to mixtures of volatile organic compounds (VOCs) emitted by growing cultures of Trichoderma from 20 strains, representing 11 different Trichoderma species.

          Results

          We identified nine Trichoderma strains that produced plant growth promoting VOCs. Exposure to mixtures of VOCs emitted by these strains increased plant biomass (37.1–41.6 %) and chlorophyll content (82.5–89.3 %). Trichoderma volatile-mediated changes in plant growth were strain- and species-specific. VOCs emitted by T. pseudokoningii (CBS 130756) were associated with the greatest Arabidopsis growth promotion. One strain, T. atroviride (CBS 01-209), in our screen decreased growth (50.5 %) and chlorophyll production (13.1 %). Similarly, tomatoes exposed to VOCs from T. viride (BBA 70239) showed a significant increase in plant biomass (>99 %), larger plant size, and significant development of lateral roots. We also observed that the tomato plant growths were dependent on the duration of the volatile exposure. A GC–MS analysis of VOCs from Trichoderma strains identified more than 141 unique compounds including several unknown sesquiterpenes, diterpenes, and tetraterpenes.

          Conclusions

          Plants grown in the presence of fungal VOCs emitted by different species and strains of Trichoderma exhibited a range of effects. This study demonstrates that the blend of volatiles produced by actively growing fungi and volatile exposure time in plant development both influence the outcome of volatile-mediated interactions. Only some of our growth promoting strains produced microbial VOCs known to enhance plant growth. Compounds such as 6-pentyl-2 H-pyran-2-one were not common to all promoting strains. We found that biostimulatory strains tended to have a larger number of complex terpenes which may explain the variation in growth induced by different Trichoderma strains.

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          Most cited references53

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          Rhizosphere bacteria help plants tolerate abiotic stress.

          Plant-growth-promoting rhizobacteria (PGPR) are associated with plant roots and augment plant productivity and immunity; however, recent work by several groups shows that PGPR also elicit so-called 'induced systemic tolerance' to salt and drought. As we discuss here, PGPR might also increase nutrient uptake from soils, thus reducing the need for fertilizers and preventing the accumulation of nitrates and phosphates in agricultural soils. A reduction in fertilizer use would lessen the effects of water contamination from fertilizer run-off and lead to savings for farmers.
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            Biosynthesis, function and metabolic engineering of plant volatile organic compounds.

            Plants synthesize an amazing diversity of volatile organic compounds (VOCs) that facilitate interactions with their environment, from attracting pollinators and seed dispersers to protecting themselves from pathogens, parasites and herbivores. Recent progress in -omics technologies resulted in the isolation of genes encoding enzymes responsible for the biosynthesis of many volatiles and contributed to our understanding of regulatory mechanisms involved in VOC formation. In this review, we largely focus on the biosynthesis and regulation of plant volatiles, the involvement of floral volatiles in plant reproduction as well as their contribution to plant biodiversity and applications in agriculture via crop-pollinator interactions. In addition, metabolic engineering approaches for both the improvement of plant defense and pollinator attraction are discussed in light of methodological constraints and ecological complications that limit the transition of crops with modified volatile profiles from research laboratories to real-world implementation. © 2013 The Authors. New Phytologist © 2013 New Phytologist Trust.
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              Biology and biotechnology of Trichoderma

              Fungi of the genus Trichoderma are soilborne, green-spored ascomycetes that can be found all over the world. They have been studied with respect to various characteristics and applications and are known as successful colonizers of their habitats, efficiently fighting their competitors. Once established, they launch their potent degradative machinery for decomposition of the often heterogeneous substrate at hand. Therefore, distribution and phylogeny, defense mechanisms, beneficial as well as deleterious interaction with hosts, enzyme production and secretion, sexual development, and response to environmental conditions such as nutrients and light have been studied in great detail with many species of this genus, thus rendering Trichoderma one of the best studied fungi with the genome of three species currently available. Efficient biocontrol strains of the genus are being developed as promising biological fungicides, and their weaponry for this function also includes secondary metabolites with potential applications as novel antibiotics. The cellulases produced by Trichoderma reesei, the biotechnological workhorse of the genus, are important industrial products, especially with respect to production of second generation biofuels from cellulosic waste. Genetic engineering not only led to significant improvements in industrial processes but also to intriguing insights into the biology of these fungi and is now complemented by the availability of a sexual cycle in T. reesei/Hypocrea jecorina, which significantly facilitates both industrial and basic research. This review aims to give a broad overview on the qualities and versatility of the best studied Trichoderma species and to highlight intriguing findings as well as promising applications.
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                Author and article information

                Contributors
                (848) 932-6235 , Samantha.LeexIII@gmail.com
                mxy5151@psu.edu
                gregbehringer@gmail.com
                RichardHung8888@gmail.com
                profmycogirl@yahoo.com
                Journal
                Fungal Biol Biotechnol
                Fungal Biol Biotechnol
                Fungal Biology and Biotechnology
                BioMed Central (London )
                2054-3085
                29 September 2016
                29 September 2016
                2016
                : 3
                : 7
                Affiliations
                [1 ]GRID grid.430387.b, ISNI 0000000419368796, Department of Plant Biology and Pathology, , Rutgers, The State University of New Jersey, ; New Brunswick, NJ USA
                [2 ]GRID grid.29857.31, ISNI 0000000120974281, Department of Biochemistry, Microbiology and Molecular Biology, , Pennsylvania State University, ; 309 Life Sciences Building, State College, PA 16803 USA
                [3 ]GRID grid.440573.1, Chemistry Program, Division of Science, , New York University Abu Dhabi, ; PO Box 129188, Abu Dhabi, United Arab Emirates
                [4 ]GRID grid.258471.d, ISNI 0000000105130152, Biology Department, , Kean University, ; 1000 Morris Ave., Union, NJ 07083 USA
                Author information
                http://orcid.org/0000-0003-0688-0048
                Article
                25
                10.1186/s40694-016-0025-7
                5611631
                28955466
                78c2a48f-6e73-4df9-a76a-66c9664d94f6
                © The Author(s) 2016

                Open AccessThis article is distributed under the terms of the Creative Commons Attribution 4.0 International License ( http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The Creative Commons Public Domain Dedication waiver ( http://creativecommons.org/publicdomain/zero/1.0/) applies to the data made available in this article, unless otherwise stated.

                History
                : 14 July 2016
                : 22 September 2016
                Funding
                Funded by: FundRef http://dx.doi.org/10.13039/100000001, National Science Foundation;
                Award ID: 0937373
                Award Recipient :
                Funded by: FundRef http://dx.doi.org/10.13039/100006102, Northeast SARE;
                Award ID: GNE14-084-27806
                Award Recipient :
                Categories
                Research
                Custom metadata
                © The Author(s) 2016

                volatile organic compounds,trichoderma,plant growth,arabidopsis thaliana,solanum lycopersicum,gas chromatography–mass spectrometry,plant–microbe interactions

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