Enhanced ethylene production and leaf epinasty are characteristic responses of tomato (Lycopersicon escukntum Mill.) to waterlogging. It has been proposed (Bradford, Yang 1980 Plant Physiol 65: 322-326) that this results from the synthesis of the immediate precursor of ethylene, 1-aminocyclopropane-I-carboxylic acid (ACC), in the waterlogged roots, its export in the transpiration stream to the shoot, and its rapid conversion to ethylene. Inhibitors of the ethylene biosynthetic pathway are available for further testing of this ACC transport hypothesis: aminooxyacetic acid (AOA) or aminoethoxyvinylglycine (AVG) block the synthesis of ACC, whereas CO2 prevents its conversion to ethylene. AOA and AVG, supplied in the nutrient solution, were found to inhibit the synthesis and export of ACC from anaerobic roots, whereas Co2" had no effect, as predicted from their respective sites of action. Transport of the inhibitors to the shoot was demonstrated by their ability to block wound ethylene synthesis in excised petioles. Al three inhibitors reduced petiolar ethylene production and epinasty in anaerobically stressed tomato plants. With AOA and AVG, this was due to the prevention of ACC import from the roots as well as inhibition of ACC synthesis in the petioles. With Co21, conversion of both root-and petiole-synthesized ACC to ethylene was blocked. Collectively, these data support the hypothesis that the export of ACC from low 02 roots to the shoot is an important factor in the ethylene physiology of waterlogged tomato plants.Ethylene is an important endogenous regulator of plant responses to waterlogging (7,14). Under the low 02 conditions in the soil during flooding, tomato roots synthesize ACC,3 the immediate precursor of ethylene, and transport it in the xylem to the shoot, where it is rapidly converted to ethylene (8,9). This increase in the ethylene content of the shoot tissues results in petiole epinasty, aerenchyma formation, and other anatomical and morphological responses associated with waterlogging (6,11,14).The sites of inhibition in the biosynthetic pathway of ethylene are now known for several inhibitors. Application of these inhibitors allows further tests of the role of root-synthesized ACC in shoot responses to waterlogging. AOA and AVG effectively prevent the conversion of SAM to ACC, both in vivo (1,22,23) conversion of ACC to ethlene. Co2" on the other hand, has no effect on ACC synthesis but inhibits the formation of ethylene from ACC (23). These compounds, therefore, permit inhibition of ethylene biosynthesis at specific steps in the pathway (see 18, 19, for reviews). In waterlogged plants, the locations of stress-induced ACC synthesis and its conversion to ethylene are separated spatially, with the formation of ACC from SAM occurring in the roots and conversion of ACC to ethylene occurring in the shoot.Thus, the effects of application of AOA, AVG, or Co2" to anaerobic roots should reflect their differing sites of action. In this study, these inhibitors were used to further characterize th...