Changes in the concentration of organic acids in roots and leaves of carob-tree under Fe deficiency

Several fruit trees are able to cope with iron (Fe) deficiency when grown in calcareous soils in the Mediterranean region, although information regarding well adapted slow-growing species is scarce, and the mechanisms activated by these species are not described in the literature. A crucial issue related to tolerance is the need to transport Fe over relatively long distances inside the plant. To evaluate the possible role of organic acids in the movement of Fe in tolerant plants, we studied the concentration of low molecular weight organic acids in several organs of 1-year old carob plants grown for 55 days in nutrient solutions without Fe (0µM Fe) or with 1µM Fe and 10µM Fe. Roots, stems and leaves were harvested, and the biomass, Fe and organic acid contents quantified. Total leaf chlorophyll (Chl) was evaluated in young leaves over the experimental period and the activity of root ferric chelate-reductase (FC-R; EC 1.16.1.17) was determined after 35 days, when deficiency symptoms appeared. Iron chlorosis was observed only at the end of the experiment in plants grown in the absence of Fe, and these plants had a smaller DW of leaves and also significant greater activity of root FC-R. Iron deficiency (Fe0 and Fe1 treatments) induced significant changes in the concentrations of succinic, malic, citric and fumaric acids, which increased in roots, or in basal, middle and apical leaves. There were significant correlations between most organic acids (with the exceptions of 2-oxoglutaric and tartaric acids) and leaf Chl. Analysis of each type of leaf showed that more succinic and malic acids were present in young chlorotic leaves while the reverse was true for quinic acid. These changes in organic acids followed a root-to-foliage pathway that was similar in all leaf types and particularly evident in young chlorotic leaves. We hypothesised that it was associated with Fe transport from roots to aboveground tissues, as there were significant differences in Fe contents between treatments with and without Fe.