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Suppressive Effects of Compost Tea on Phytopathogens 247below 10 –18 M <strong>in</strong> soil solutions at neutral pH (Handelsman and Stabb, 1996).Iron salts that have low solubility such as iron oxides, carbonates, phosphates,hydroxides and some forms of <strong>in</strong>soluble chelates are created <strong>in</strong> certa<strong>in</strong> soiltypes that make iron not readily available to <strong>plant</strong>s. To circumvent the solubilityproblem, many microorganisms synthesize and utilize very specificlow molecular weight (500–1000 Da) iron chelators called siderophores (thatcan b<strong>in</strong>d iron and are sent out by the cells to absorb iron, as well as be<strong>in</strong>g used<strong>in</strong>side the cell to store it). When grown under iron-deficient conditions, manymicrobes will synthesize and excrete siderophores <strong>in</strong> excess of their own drycell weight to kidnap and solubilize iron and made it unavailable to the pathogens.The typical system <strong>in</strong>volves a siderophore, which is an iron-b<strong>in</strong>d<strong>in</strong>gligand, and an uptake prote<strong>in</strong>, which transports the siderophore <strong>in</strong>to the cell.The fluorescent pseudomonads produce a class of siderophore known as thepseudobact<strong>in</strong>s, which are structurally complex iron-b<strong>in</strong>d<strong>in</strong>g molecules. Analysesof mutants lack<strong>in</strong>g the ability to produce siderophores suggest that theycontribute to the suppression of certa<strong>in</strong> fungal and oomycete diseases (Duijffet al., 1994; Buysens et al., 1996). An <strong>in</strong>terest<strong>in</strong>g aspect of siderophore biologyis that diverse organisms can use the same type of siderophore. Microorganismsmay use each other’s siderophores if they conta<strong>in</strong> the appropriate uptakeprote<strong>in</strong> (Koster et al., 1993; Raaijmakers et al., 1995), and <strong>plant</strong>s can evenacquire iron from certa<strong>in</strong> pseudobact<strong>in</strong>s (Duijff et al., 1994).Microorganisms <strong>in</strong> compost produce siderophores that keep iron <strong>in</strong> anavailable form for <strong>plant</strong>s <strong>in</strong> the soil, even at high pH. Compost also produceswater-soluble humic substances, <strong>in</strong>clud<strong>in</strong>g fulvic acids, which keep iron, z<strong>in</strong>c,manganese and other trace elements <strong>in</strong> solution (Chen and Inbar, 1993).AntibiosisAntibiosis is def<strong>in</strong>ed as antagonism mediated by specific or non-specificmetabolites of microbial orig<strong>in</strong>, by lytic agents, enzymes, volatile compoundsor other toxic substances (Jackson, 1965; Fravel, 1988). Antibiotic productionappears to be important to the survival of microorganisms through the elim<strong>in</strong>ationof microbial competition for food sources, which are usually verylimited <strong>in</strong> soil (Ellis et al., 2000; Slattery et al., 2001). Antibiotic production isvery common among soil-dwell<strong>in</strong>g bacteria and fungi. Inhibition <strong>in</strong> the Petridish may be the result of antibiosis, but it is not easy to show that this antibiosisis actually responsible for disease suppression. First, the antibioticmust be extracted, purified and identified chemically. Then it is necessary toshow that the microorganism grows <strong>in</strong> the microhabitat of the pathogen, andthat the antibiotic is produced <strong>in</strong> the right place, at the right time, and <strong>in</strong> sufficientamounts to control disease. It is also necessary to demonstrate that thepathogen is sensitive to the antibiotic. Genetic analyses have been particularly<strong>in</strong>formative <strong>in</strong> determ<strong>in</strong><strong>in</strong>g the role of antibiotics <strong>in</strong> biocontrol, <strong>in</strong> partbecause mutants can be screened easily <strong>in</strong> vitro for changes <strong>in</strong> antibioticaccumulation, provid<strong>in</strong>g the means to conduct thorough genetic analyses(Handelsman and Stabb, 1996).

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