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11.2 Biocontrolj215<br />

graminis var. tritici but failed to suppress two other isolates and the pathogen was<br />

resistant to DAPG at 3 enzyme units per ml. DAPG-producing fluorescent Pseudomonas<br />

spp. have been shown to be responsible for take-all decline, a natural biological<br />

control system found to develop in soils following extended monoculture of<br />

wheat or barley [29,30]. It is possible to isolate DAPG from the rhizosphere. Therefore,<br />

the positive role of DAPG in the biological control of plant disease can be<br />

assessed <strong>by</strong> genetic approaches. Raaijmakers et al. [31] have demonstrated that the<br />

level of DAPG in the rhizosphere is directly related to the DAPG-producing population.<br />

Phloroglucinol (Phl) is a phenolic metabolite produced <strong>by</strong> bacteria and plants<br />

with broad-spectrum antibacterial, antifungal, antiviral, antihelmintic and phytotoxic<br />

properties [32]. This polyketide antibiotic has been identified to be largely responsible<br />

for the prevention of damping-off in sugarbeet and cotton caused <strong>by</strong> Pythium<br />

ultimum and Phytium spp., respectively [27,33].<br />

The biocontrol strain P. fluorescens F113 is an effective antagonist of Pythium<br />

ultimum under laboratory conditions [27] besides reducing the severity of dampingoff<br />

in soil naturally infested with Pythium spp. [34]. P. fluorescens F113G22, a Phlnegative<br />

Tn5::lacZY mutant derivative, does not inhibit P. ultimum grown in vitro or<br />

reduce the severity of damping-off [27]. The Phl biosynthetic locus has been cloned<br />

in several pseudomonads [34–40]. In microcosm studies, these two Phl overproducing<br />

strains proved to be as effective in controlling damping-off disease as a proprietary<br />

fungicide treatment, indicating enhanced potential of genetic modification in<br />

plant disease control [41].<br />

Pyoluteorin, an aromatic polyketide antibiotic, is produced <strong>by</strong> several Pseudomonas<br />

species including strains that suppress plant diseases caused <strong>by</strong> phytopathogenic<br />

fungi [33,42,43]. Howell and Stipanovic [33] reported that pyoluteorin<br />

treatment was effective in providing protection against damping-off caused <strong>by</strong><br />

Pythium. Of the antibiotics known to be produced <strong>by</strong> Pseudomonas fluorescens<br />

Pf-5 or CHA0, pyoluteorin is most toxic to Pythium ultimum [42], although<br />

2,4-diacetylphloroglucinol [44,45] and pyoverdine siderophores [46] also suppress<br />

mycelial growth [33].<br />

Howell and Stipanovic [47] reported that pyrrolnitrin plays an important role in<br />

providing protection against Rhizoctonia solani infection in cotton seedlings. Several<br />

studies suggest that pyrrolnitrin production <strong>by</strong> Burkholderia cepacia and Pseudomonas<br />

spp. is closely related to biocontrol of plant diseases. Jayaswal et al. [48,49]<br />

generated a Tn5-induced mutant strain of B. cepacia deficient in pyrrolnitrin and<br />

showed that the mutant completely lost antifungal activity. Furthermore, Hill et al.<br />

[50] cloned a gene responsible for pyrrolnitrin production in P. fluorescens and<br />

demonstrated that interruption within the gene region resulted in the loss of biocontrol<br />

activity against Rhizoctonia damping-off in cotton. A chemically induced<br />

overproducing mutant of P. aeruginosa exhibited 30-fold increase in synthesis of<br />

pyrrolnitrin [51]. Replacing the native promoter with a more active promoter within<br />

Prn gene cluster also increased pyrrolnitrin production in P. aureginosa with<br />

enhanced biocontrol of Rhizoctonia damping-off [52].<br />

1-Aminocyclopropane-1-carboxylic acid (ACC) deaminase, which is found only in<br />

microorganisms, catalyzes cleavage of ACC to a-ketobutyrate and ammonia <strong>by</strong>

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