Phloem unloading following reactivation in predarkened mature maize leaves

Abstract

Mature leaf blades of 48-h predarkened maize plants (Zea mays L. cv. Prior) were excised, and treated apically as the source (light, normal air) and basally as the sink (light or dark, air without CO₂). After providing the source portion with ¹⁴CO₂, the sink portions were harvested after 2, 7 or 14 h by freezing with liquid nitrogen, grinding, and freeze-drying. Extracts, fractionated by ionexchange resins into neutral, basic and acid fractions, were chromatographed on thin cellulose layers, and autoradiographed. Identification of labeled compounds was carried out by co-chromatography with authentic labeled substances. Activities of enzymes pertaining to the metabolism of sucrose were checked. Results show that the source supplies sucrose to the sink, where it is unloaded and metabolized by acid invertase (EC 3.2.1.26) in both the light and the dark. Starch appearing in the sink only in the light, after 7 h of re-illumination, yields labeled glucose upon hydrolysis. Although sucrose-phosphate synthetase (EC 2.4.1.14) is active in sinks and in isolated vascular-bundle fragments, it remains questionable whether sucrose unloaded from sieve tubes is metabolized by a method other than inversion. Sucrose synthetase (EC 2.4.1.13) was found to be inactive. Obviously, the main metabolite of unloaded sucrose is glucose-6-phosphate, giving access to the glycolytic pathway. The main difference between the sinks in the light and the dark is the lack of labeled glycine and serine in the dark. This indicates that in the light decarboxylation of glycine yields CO₂, which is recycled photosynthetically.