Seasonal dynamics of mineral nutrients and carbohydrates by walnut tree leaves

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Abstract
The dry weight accumulation per leaf as well as the concentration per gram of dry weight and the accumulation of nitrogen (N), phosphorus (P), potassium (K), calcium (Ca), magnesium (Mg), iron (Fe), manganese (Mn), zinc (Zn), and copper (Cu) were determined in walnut tree leaves (Juglans regia L.) during a complete life cycle. Additionally, the dynamics of plant nutrient concentration in leaf petiole sap and carbohydrate accumulation in leaves were studied in relation to the main life cycle events of the walnut tree. Total N, P, K, Cu, and Zn concentrations decreased, whereas that of Ca, Mg, and Mn increased during the season. Iron concentration fluctuated around a mean value. Total N, P, K, Mg, and Cu concentrations detected in younger mature leaves were at the sufficient level, whereas Ca, Fe, Mn, and Zn concentrations were at higher levels as compared to those previously reported. All the detected nutrient accumulations increased abruptly during leaf ontogeny and leaf maturation until a maximum level was attained in the younger mature leaves. Similarly, sucrose, glucose, and fructose accumulation were observed at the same period. The rates of total N, P, Cu, and Zn accumulation were lower than the rates of the observed dry matter accumulation and nutrient concentration dilution. Potassium and Mn accumulation rates were almost equal, whereas those for Ca and Mg were higher as compared to the dry matter accumulation rate. The fast embryo growing phase resulted in a considerable decrease in dry weight, total N, P, K, Cu, Zn, and carbohydrate accumulation, and to a lesser degree in Ca, Mg, and Mn accumulation. Nutrient accumulation reduction in leaves by the influence of the growing fruits were estimated to be: total N 52%, K 48%, P 29.5%, Mg 16.3%, Ca 15%, Fe 51.2%, Cu 55.2%, Zn 37.3%, and Mn 5.4% of the maximum nutrient value of the younger mature leaves. Old leaves preserved nutrients before leaf fall as follows: total N 25.4%, P 45%, K 31%, Ca 74.8%, Mg 76.5%, Mn 89.2%, Fe and Zn 50%, and Cu 37%. Nutrient remobilization from the senescing old leaves before leaf fall were: total N 22.6%, P 25.5%, K 21%, Ca 10.2%, Mg 7%, Fe 3.2%, Mn 5.4%, Cu 8%, and Zn 13.3% of the maximum value in the younger mature leaves. In early spring, the absorption rates of N, P, and Ca were low while those of Mg, Fe, Mn, Cu, and Zn were high. During the fast growing pollen phase, the N, P, Fe, Mn, Cu, and Zn concentrations were reduced. Calcium concentration is supposed to be more affected by the rate of transpiration rather than during the growing of embryo. Calcium and Mg concentrations in the sap were negatively correlated. The detected K concentration level in the sap was as high as 33 to 50 times that of soluble N, 12 to 21 times to that of P, 5 times to that of Ca, and 10 to 20 times to that of Mg. The first maximum of starch accumulation in mature leaves was observed during the slow growing embryo phase and a second one after fruit ripening. Old senescing leaves showed an extensive carbohydrate depletion before leaf fall.