Since the discovery of the plant growth-regulating properties of TRIA, a primary alcohol that is a natural constituent of plant waxes, and its second messenger L(+)-adenosine, the rapid response kinetics to these compounds have been enigmatic (Ries and Wert, 1988;Ries, 1991).TRIA increased the dry weight, free amino acids, reducing sugars, and soluble protein of rice (Oryza sativa L.) and maize (Zea mays L.) plants within 5 min (Ries, 1991). TRIA also elicited the appearance of L(+)-adenosine in the roots of plants whose shoots were sprayed with nanomolar concentrations within 1 min (Ries and Wert, 1988). This was the first evidence that L(+)-adenosine occurred in nature. Synthetic L(+)-adenosine increased the rate of growth of rice seedlings, as measured by total dry weight gain, by more than 50% within 24 h of foliar application of 0.01 to 100.0 kg L-' (3.7 X 10-" to 10-7 M), whereas D(-)-adenosine did not affect plant growth (Ries, 1991).
49site plant part, providing it was applied 1 min prior to TRIA application (Ries and Wert, 1988). TRIA applied to oat (Avena sativa L.) or tomato (Lycopersicon esculentum Mill.) shoots connected to rice roots by a 4-mm water column also resulted in the appearance of L(+)-adenosine (TRIM) in rice roots (Ries and Wert, 1988).In an attempt to determine other physiological responses to TRIA in addition to the elicitation of L(+)-adenosine, 20-to 25-d-old tomato seedlings were sprayed with water or TRIA, and after 1 min the plants were excised. Analysis of the diffusate from the excised shoots, as determined by HPLC and atomic absorption spectrophotometry, indicated large concentration differences in organic compounds and inorganic cations (unpublished data). The largest differences were in the cation concentration of the exudate from the stump of the excised tomato plant. Thus, the objective of this research was to use this observation to further elucidate the mode of action for the rapid responses of plants to TRIA and L(+)-adenosine.We present here evidence that foliar applications of both of these compounds at nanomolar concentrations cause rapid changes in soluble Ca2+, Mg2+, and K+ concentrations within xylem exudates from the stumps of excised stems and leaves.
MATERIALS A N D METHODS
Plant Crowth and TreatmentTomatoes (Lycopersicon esculentum Mill. cv Sunny), cucumbers (Cucumis sativa L. cv Flurry), and maize (Zea mays L.cv Pioneer 3780) were grown in a greenhouse with approximately 16 h of supplemental light (700 pmol s-' m-', metal halide) daily. Seeds were planted in 15-cm diameter clay pots containing a soil mix, and the plants were thinned to two or three per pot 8 to 10 d after emergence. Soluble fertilizer (20 N-8.6 P-16.6 K; 1.0 g L-' Peters 20-20-20, W.R. Grace and Co., Fogelsville, PA) was applied once or twice after planting and again prior to treatment. The pots were labeled, randomized for treatments within blocks, and isolated from each other on the greenhouse bench. They were not disturbed for several hours prior to initiation of the treatments.Experiment...