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ALLELOPATHIC BACTERIA IN WEED MANAGEMENT 147<br />

Successful competition of bacteria living in the rhizosphere depends on several<br />

factors, including rapid growth on multiple substrates, antibiotic production, and<br />

downward growth with the root. A major factor contributing to successful competition<br />

of rhizobacteria over other microorganisms is the growth stimulation by exuded organic<br />

compounds and sloughed-off root hair and epidermal cell materials (De Weger et al.,<br />

1995). The ability to efficiently compete for these available resources and to produce<br />

siderophores for obtaining iron is important in establishment, colonization, and<br />

persistence of rhizobacteria in the rhizosphere.<br />

A characteristic of many AB is the high specificity toward their weed host(s)<br />

with no detrimental effects on growth of nonweedy plant species (Cherrington and<br />

Elliott, 1987; Elliott and Lynch, 1985; Kennedy et al., 1991; 2001). Although effects<br />

on plants are subtle (Kremer and Kennedy, 1996), AB may be as significant as<br />

traditional bacterial pathogens in affecting plant growth (Schroth and Hancock, 1982;<br />

Suslow and Schroth, 1982). Because AB attack the seed and/or seedling rather than<br />

the growing plant, weed seed or vegetative propagule production is suppressed, a key<br />

to any weed management program, which reduces the need for repeated postemergence<br />

herbicide applications and increases the chances of success for control of a growing,<br />

competitive weed (Aldrich and Kremer, 1997).<br />

4. MODES OF ACTION OF ALLELOPATHIC BACTERIA<br />

Many AB strains produce secondary metabolites that are inhibitory to plants, including<br />

phytotoxins and antibiotics, which can be considered allelopathic. Phytotoxins from<br />

fluorescent Pseudomonas spp., a diverse group of plant pathogenic bacteria abundant<br />

in the soil and rhizosphere, have been well studied (Mitchell, 1991). There are fewer<br />

reports on phytotoxins from AB and many have not been extensively studied.<br />

A phytotoxin from Pseudomonas fluorescens strain D7 was shown to be<br />

responsible for root growth inhibition of downy brome (Bromus tectorum) (Tranel et<br />

al., 1993). Further characterization revealed that the active fraction was a complex of<br />

chromopeptides, other peptides and fatty acid esters in a lipopolysaccharide matrix<br />

(Gurusiddaiah et al., 1994). Secondary metabolites isolated from Pseudomonas<br />

syringae strain 3366 inhibitory to downy brome consisted of phenazine-1-carboxylic<br />

acid, 2-aminophenoxazone and 2-aminophenol (Gealy et al., 1996). Gealy et al.<br />

(1996) showed that phenazine-type antibiotics of Pseudomonas fluorescens also<br />

inhibited downy brome root growth. Electron microscopy of AB colonizing the<br />

rhizoplane and endorhizal cells of leafy spurge (Euphorbia esula) revealed disruption<br />

of plant cell walls and membranes apparently due to production of phytotoxins and/or<br />

enzymes by the bacteria, which consequently inhibited seedling growth (Souissi et<br />

al., 1997). AB may also produce “phytotoxic antibiotics” that affect plant growth<br />

such as the broad-spectrum antibiotic, 2,4-diacetylphloroglucinol, released by P.<br />

fluorescens strain CHA0, which suppressed soilborne fungal plant pathogens but was<br />

also highly phytotoxic to seedlings of several plant species (Keel et al., 1992).<br />

Plant-inhibitory effects of some AB are auxin-mediated, illustrated by direct uptake<br />

of bacterially produced indoleacetic acid (IAA). Plant response to microbially

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