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8 ACC Deaminase-Containing Plant Growth-Promoting Bacteria 137<br />

promoting bacteria, Pseudomonas putida ATCC 17399 and Pseudomonas fluorescens<br />

ATCC 17400, by conjugational transfer, the transconjugants acquired<br />

the ability to grow on minimal media using ACC as the sole source of nitrogen,<br />

and to promote the elongation of canola roots (Shah et al. 1998).<br />

In 1998, Glick et al. proposed a model in which <strong>plant</strong> growth-promoting<br />

bacteria can lower <strong>plant</strong> ethylene levels and in turn stimulate <strong>plant</strong> growth. In<br />

this model, the <strong>plant</strong> growth-promoting bacteria bind to the <strong>surface</strong> of either<br />

the seed or root of a developing <strong>plant</strong>; in response to tryptophan and other<br />

small molecules in the seed or root exudates (Whipps 1990), the <strong>plant</strong> growthpromoting<br />

bacteria synthesize and secrete indole acetic acid (IAA; Fallik et al.<br />

1994; Patten and Glick 1996), some of which is taken up by the <strong>plant</strong>. This IAA<br />

together with endogenous <strong>plant</strong> IAA, can stimulate <strong>plant</strong> cell proliferation,<br />

<strong>plant</strong> cell elongation or induce the activity of ACC synthase to convert SAM to<br />

ACC (Kende 1993).<br />

Much of the ACC produced by this latter reaction is exuded from seeds or<br />

<strong>plant</strong> roots along with other small molecules normally present in seed or root<br />

exudates (Penrose and Glick 2001). The ACC in the exudates may be taken up<br />

by the bacteria and subsequently hydrolyzed by the enzyme, ACC deaminase,<br />

to ammonia and a-ketobutyrate. The uptake and cleavage of ACC by <strong>plant</strong><br />

growth-promoting bacteria decreases the amount of ACC outside the <strong>plant</strong>.<br />

Increasing amounts of ACC are exuded by the <strong>plant</strong> in order to maintain the<br />

equilibrium between internal and external ACC levels.As a result of the activity<br />

of ACC deaminase, the presence of the bacteria induces the <strong>plant</strong> to synthesize<br />

more ACC than it would otherwise need and as well, stimulates the<br />

exudation of ACC from the <strong>plant</strong>.<br />

Thus, <strong>plant</strong> growth-promoting bacteria are supplied with a unique source<br />

of nitrogen in the form of ACC that enables them to proliferate under conditions<br />

in which other soil bacteria may not flourish. As a result of lowering the<br />

ACC level within the <strong>plant</strong>, either the endogenous level or the IAA-stimulated<br />

level, the amount of ethylene in the <strong>plant</strong> is also reduced.<br />

Plant growth-promoting bacteria that possess the enzyme ACC deaminase<br />

and are bound to seeds or roots of seedlings, can reduce the amount of <strong>plant</strong><br />

ethylene and the extent of its inhibition on root elongation. Thus, these <strong>plant</strong>s<br />

should have longer roots and possibly longer shoots as well, inasmuch as stem<br />

elongation is also inhibited by ethylene, except in flooding-resistant <strong>plant</strong>s<br />

(Abeles et al. 1992).<br />

3.1 Treatment of Plants with ACC Deaminase Containing Bacteria<br />

Consistent with the above mentioned model, ACC deaminase activity was<br />

completely lost and the ability to promote the elongation of canola roots<br />

under gnotobiotic conditions was greatly diminished when the ACC deaminase<br />

gene (acdS) from Enterobacter cloacae UW4 was replaced, by homolo-

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