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134<br />

Bernard R. Glick and Donna M. Penrose<br />

bacteria may vary from one species of <strong>plant</strong> to another, as well as according to<br />

the age of the <strong>plant</strong>.<br />

2 Ethylene<br />

In higher <strong>plant</strong>s ethylene is produced from L-methionine via the intermediates,<br />

S-adenosyl-L-methionine (SAM) and 1-aminocyclopropane-1-carboxylic<br />

acid (ACC; Yang and Hoffman 1984). The enzymes involved in this<br />

metabolic sequence are SAM synthetase, which catalyzes the conversion of<br />

methionine to SAM (Giovanelli et al. 1980); ACC synthase, which is responsible<br />

for the hydrolysis of SAM to ACC and 5¢–methylthioadenosine (Kende<br />

1989) and ACC oxidase which further metabolizes ACC to ethylene, carbon<br />

dioxide, and cyanide (John 1991).<br />

Ethylene, which is produced in almost all <strong>plant</strong>s, mediates a range of <strong>plant</strong><br />

responses and developmental steps. Ethylene is involved in seed germination,<br />

tissue differentiation, formation of root and shoot primordia, root elongation,<br />

lateral bud development, flowering initiation, anthocyanin synthesis, flower<br />

opening and senescence, fruit ripening and degreening, production of volatile<br />

organic compounds responsible for aroma formation in fruits, storage product<br />

hydrolysis, leaf and fruit abscission, and the response of <strong>plant</strong>s to biotic<br />

and abiotic stress (Matoo and Suttle 1991; Abeles et al. 1992; Frankenberger<br />

and Arshad 1995). In some instances, ethylene is stimulatory while in others it<br />

is inhibitory.<br />

The term “stress ethylene” was coined by Abeles (1973) to describe the<br />

acceleration of ethylene biosynthesis associated with biological and environmental<br />

stresses, and pathogen attack (Morgan and Drew 1997). The increased<br />

level of ethylene formed in response to trauma inflicted by chemicals, temperature<br />

extremes, water stress, ultraviolet light, insect damage, disease, and<br />

mechanical wounding (Bestwick and Ferro 1998) can be both the cause of<br />

some of the symptoms of stress (e.g., onset of epinastic curvature and formation<br />

of arenchyma), and the inducer of responses which will enhance survival<br />

of the <strong>plant</strong> under adverse conditions (e.g., production of antibiotic enzymes<br />

and phytoalexins).<br />

Chemicals have been used to control ethylene levels in <strong>plant</strong>s. The application<br />

of compounds such as rhizobitoxin, an amino acid secreted by several<br />

strains of bacteria, and its synthetic analog, aminoethoxyvinylglycine (AVG),<br />

can inhibit ethylene biosynthesis; silver thiosulfate can inhibit ethylene action,<br />

and 2-chloroethylphosphoric acid (ethephon), regarded by some researchers<br />

as “liquid ethylene”, can release ethylene (Abeles et al. 1992). Sisler and Serek<br />

(1997) discovered that cyclopropenes can block ethylene perception and are<br />

potentially useful for extending the life span of cut flowers and the display life<br />

of potted <strong>plant</strong>s. In addition, tropolone compounds were isolated from wood<br />

by Mizutani et al. (1998). These compounds, which can inhibit the growth of

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