Storage ability of overwintering leaves and rhizomes in a semi-evergreen fern, Dryopteris crassirhizoma (Dryopteridaceae)

Abstract

Dryopteris crassirhizoma is a rhizomatous semi-evergreen fern growing in the understory of deciduous forests. Although the top portion of the overwintering leaves began to wither in early winter, intensive senescence occurred in the spring, concurrently with new leaf development. Dry weight comparisons between organs revealed that the rhizome occupied the largest proportion of the total mass, followed by the pinnae. To assess the storage ability of overwintering leaves and the rhizome, seasonal changes in nitrogen content and the dry mass of pinnae and the rhizome were measured. Nitrogen (36.6%) was resorbed from winter-withering pinnae, but not from spring-withering pinnae. In contrast, a similar decrease in dry mass per unit area occurred between winter- and springwithering pinnae (∼15%). These results indicate that overwintering leaves serve as a carbohydrate storage organ, but do not serve as a nitrogen storage organ. Nitrogen was not translocated from the rhizome during the early growing season, but translocation did occur in late summer and autumn. The dry mass of the rhizome decreased by 18.4% in spring, at the time of new leaf expansion. The amount of exported dry matter from the rhizome was threefold larger than that from senescent pinnae. Therefore, the rhizome is a major carbohydrate storage organ in this species, although overwintering leaves also act as a carbohydrate storage organ.