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Taxonomy and phylogeny of Gliocladium analysed from nuclear large subunit ribosomal DNA sequences

Microbial biological control agents (MBCAs) are applied to crops for biological control of plant pathogens where they act via a range of modes of action. Some MBCAs interact with plants by inducing resistance or priming plants without any direct interaction with the targeted pathogen. Other MBCAs act via nutrient competition or other mechanisms modulating the growth conditions for the pathogen. Antagonists acting through hyperparasitism and antibiosis are directly interfering with the pathogen. Such interactions are highly regulated cascades of metabolic events, often combining different modes of action. Compounds involved such as signaling compounds, enzymes and other interfering metabolites are produced in situ at low concentrations during interaction. The potential of microorganisms to produce such a compound in vitro does not necessarily correlate with their in situ antagonism. Understanding the mode of action of MBCAs is essential to achieve optimum disease control. Also understanding the mode of action is important to be able to characterize possible risks for humans or the environment and risks for resistance development against the MBCA. Preferences for certain modes of action for an envisaged application of a MBCA also have impact on the screening methods used to select new microbials. Screening of MBCAs in bioassays on plants or plant tissues has the advantage that MBCAs with multiple modes of action and their combinations potentially can be detected whereas simplified assays on nutrient media strongly bias the selection toward in vitro production of antimicrobial metabolites which may not be responsible for in situ antagonism. Risks assessments for MBCAs are relevant if they contain antimicrobial metabolites at effective concentration in the product. However, in most cases antimicrobial metabolites are produced by antagonists directly on the spot where the targeted organism is harmful. Such ubiquitous metabolites involved in natural, complex, highly regulated interactions between microbial cells and/or plants are not relevant for risk assessments. Currently, risks of microbial metabolites involved in antagonistic modes of action are often assessed similar to assessments of single molecule fungicides. The nature of the mode of action of antagonists requires a rethinking of data requirements for the registration of MBCAs.

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Mode of Action of Microbial Biological Control Agents Against Plant Diseases: Relevance Beyond Efficacy

Biocontrol involves harnessing disease-suppressive microor? ganisms to improve plant health. Disease suppression by biocontrol agents is the sustained manifestation of interactions among the plant, the pathogen, the biocontrol agent, the microbial community on and around the plant, and the physi? cal environment. Even in model laboratory systems, the study of biocontrol involves interactions among a minimum of three organisms. Therefore, despite its potential in agricultural ap? plications, biocontrol is one of the most poorly understood areas of plant-microbe interactions. The complexity of these systems has influenced the accep? tance of biocontrol as a means of controlling plant diseases in two ways. First, practical results with biocontrol have been variable. Thus, despite some stunning successes with biocon? trol agents in agriculture, there remains a general skepticism born of past failures (Cook and Baker, 1983; Weiler, 1988). Second, progress in understanding an entire system has been slow. Recently, however, substantial progress has been made in a number of biocontrol systems through the application of genetic and mathematical approaches that accommodate the complexity. Biocontrol of soilborne diseases is particularly complex be? cause these diseases occur in the dynamic environment at the interface of root and soil known as the rhizosphere, which is defined as the region surrounding a root that is affected by it. The rhizosphere is typified by rapid change, intense microbial activity, and high populations of bacteria compared with nonrhizosphere soil. Plants release metabolically active cells from their roots and deposit as much as 20% of the carbon allocated to roots in the rhizosphere, suggesting a highly evolved relationship between the plant and rhizosphere microorgan? isms. The rhizosphere is subject to dramatic changes on a short temporal scale?rain events and daytime drought can result in fluctuations in salt concentration, pH, osmotic poten? tial, water potential, and soil particle structure. Over longer temporal scales, the rhizosphere can change due to root growth, interactions with other soil biota, and weathering processes. It is the dynamic nature of the rhizosphere that makes it an interesting setting for the interactions that lead to disease and biocontrol of disease (Rovira, 1965,1969,1991; Hawes, 1991; Waisel et al., 1991).

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Biocontrol of Soilborne Plant Pathogens.

When grown in dual culture, hyphae of the majority of Trichoderma isolates coiled around hyphae of different test fungi. Penetration of hyphae by Trichoderma hyphae seldom occurred. The importance of antibiotics and extracellular enzymes during hyphal interaction is discussed.

Antagonistic properties of species-groups of Trichoderma: III. Hyphal interaction

Muscodor crispans is a recently described novel endophytic fungus of Ananas ananassoides (wild pineapple) growing in the Bolivian Amazon Basin. The fungus produces a mixture of volatile organic compounds (VOCs); some of the major components of this mixture, as determined by GC/MS, are propanoic acid, 2-methyl-, methyl ester; propanoic acid, 2-methyl-; 1-butanol, 3-methyl-;1-butanol, 3-methyl-, acetate; propanoic acid, 2-methyl-, 2-methylbutyl ester; and ethanol. The fungus does not, however, produce naphthalene or azulene derivatives as has been observed with many other members of the genus Muscodor. The mixture of VOCs produced by M. crispans cultures possesses antibiotic properties, as does an artificial mixture of a majority of the components. The VOCs of the fungus are effective against a wide range of plant pathogens, including the fungi Pythium ultimum, Phytophthora cinnamomi, Sclerotinia sclerotiorum and Mycosphaerella fijiensis (the black sigatoka pathogen of bananas), and the serious bacterial pathogen of citrus, Xanthomonas axonopodis pv. citri. In addition, the VOCs of M. crispans killed several human pathogens, including Yersinia pestis, Mycobacterium tuberculosis and Staphylococcus aureus. Artificial mixtures of the fungal VOCs were both inhibitory and lethal to a number of human and plant pathogens, including three drug-resistant strains of Mycobacterium tuberculosis. The gaseous products of Muscodor crispans potentially could prove to be beneficial in the fields of medicine, agriculture, and industry.

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Volatile antimicrobials from Muscodor crispans, a novel endophytic fungus.

Isolates from different species-groups of Trichoderma were tested for production of non-volatile antibiotics, by an agar layer technique. Preliminary studies on the chemical nature of these antibiotics were made. Many isolates produced non-volatile antibiotics active against a range of fungi. The ability to produce such antibiotics varied between isolates of the same species-group as well as between isolates of different species groups. The susceptibility of fungi to these antibiotics varied widely; Fomes annosus (Fr.) Cooke was the most susceptible and Fusarium oxysporum Schlecht, ex Fr. the most resistant of the test fungi used. Gliotoxin and viridin were not produced, but other chloroform-soluble antibiotics, including trichodermin and peptide antibiotics were detected.

J. Webster

Papers

Antagonistic properties of species-groups of Trichoderma: II. Production of volatile antibiotics