Seasonal development of phloem in Siberian larch stems

Abstract

The seasonal development of phloem in the stems of Siberian larch (Larix sibirica Ldb.) was studied over two seasons on 50-60-year-old trees growing in a natural stand in the Siberian forest-steppe zone. Trees at the age of 20-25 years were used to study metabolites in differentiating and mature phloem elements, cambial zone, and radially growing xylem cells in the period of early and late wood formation. The development of the current-year phloem in the stems of 50-60-year-old trees started, depending on climatic conditions, in the second-third decades of May, 10-20 days before the xylem formation, and ended together with the shoot growth cessation in late July. Monitoring of the seasonal activity of cambium producing phloem sieve cells and the duration of their differentiation compared to the xylem derivatives in the cambium demonstrated that the top production of phloem and xylem cells could coincide or not coincide during the season, while their differentiation activity was always in antiphase. Sieve cells in the early phloem are separated from those in the late phloem by a layer of tannin-containing cells, which are formed in the period when late xylem formation starts. The starch content in the structural elements of phloem depends on the state of annual xylem layer development. The content of low molecular weight carbohydrates, amino acids, organic acids, and phenols in phloem cells, cambial zone, and xylem derivatives of the cambium depends on the cell type and developmental stage as well as on the type of forming wood (early or late) differing by the cell wall parameters and, hence, by the requirement for assimilates. Significant differences in the dynamics of substances per dry weight and cell were observed during cell development.