Postulated Physiological Roles of the Seven-carbon Sugars, Mannoheptulose, and Perseitol in Avocado

Abstract

Avocado (Persea americana Mill.) tissues contain high levels of the seven-carbon (C7) ketosugar mannoheptulose and its polyol form, perseitol. Radiolabeling of intact leaves of `Hass' avocado on `Duke 7' rootstock indicated that both perseitol and mannoheptulose are not only primary products of photosynthetic CO2 fixation but are also exported in the phloem. In cell-free extracts from mature source leaves, formation of the C7 backbone occurred by condensation of a three-carbon metabolite (dihydroxyacetone-P) with a four-carbon metabolite (erythrose-4-P) to form sedoheptulose-1,7-bis-P, followed by isomerization to a phosphorylated d-mannoheptulose derivative. A transketolase reaction was also observed which converted five-carbon metabolites (ribose-5-P and xylulose-5-P) to form the C7 metabolite, sedoheptulose-7-P, but this compound was not metabolized further to mannoheptulose. This suggests that C7 sugars are formed from the Calvin Cycle, not oxidative pentose phosphate pathway, reactions in avocado leaves. In avocado fruit, C7 sugars were present in substantial quantities and the normal ripening processes (fruit softening, ethylene production, and climacteric respiration rise), which occurs several days after the fruit is picked, did not occur until levels of C7 sugars dropped below an apparent threshold concentration of ≈20 mg·g-1 fresh weight. The effect of picking could be mimicked by girdling the fruit stalks, which resulted in ripening on the tree. Again, ripening followed a decline in C7 sugars to below an apparent threshold level. Taken together, these data indicate that the C7 sugars play important roles in carbon allocation processes in the avocado tree, including a possible novel role as phloem-mobile ripening inhibitors.