Further Studies of the Phloem Loading Process in Leaves of Barley and Spinach. The Comparison of Metabolite Concentrations in the Apoplastic Compartment with those in the Cytosolic Compartment and in the Sieve Tubes1

Abstract The concentrations of sucrose, amino acids, nitrate and malate in the apoplastic compartment of illuminated leaves of barley and spinach were determined and compared with the corresponding concentrations in the cytosolic compartment of mesophyll cells and in the phloem sap, as measured previously with plants grown under identical conditions. The concentrations of sucrose and amino acids in the apoplast are found to be much lower than in the cytosol and in the phloem sap, indicating that not only the uptake into the phloem of sucrose, but also of amino acids, requires transport against a concentration gradient. The gradient of sucrose and amino acids between the cytosol and the apoplast was maintained when phloem transport had been blocked by cold girdling. Apparently, the efflux of sucrose and amino acids from the source cells to the apoplast is regulated in such a way that it meets the requirements of phloem transport. The percentages of the single amino acids as part of the total amino acids are quite similar in the cytosol, apoplast and phloem sap. The ratio of sucrose to the total amino acids in the cytosol is similar to that in the apoplast but about five times higher in the phloem sap. It appears from these results that the preferential extraction of sucrose over amino acids from the source cells to the phloem is due to the uptake from the apoplast into the phloem.