Development of successive cambia and formation of secondary xylem in Suaeda nudiflora and S. fruticosa (Amaranthaceae s.l.)

Abstract

Amaranthaceae s.l. and its allied families like Amaranthaceae s.s., Chenopodiaceae s.s. and Phytolaccaceae s.s. are always debated in the literature about their pattern of secondary growth; when their stem diameter increases by forming successive cambia. In the present study, development of successive cambia and differentiation of its derivatives was investigated in Suaeda fruticosa and S. nudiflora (Amaranthaceae s.l.). After primary growth, the first complete ring of cambium was formed by the successive cambium due to failure in the development of interfascicular cambium between adjacent vascular bundles. The first ring of successive cambium initiated from the parenchymatous cells (pericyclic derivatives) external to the protophloem. This newly formed cambium was functionally bidirectional and produced secondary xylem centripetally and phloem centrifugally. Subsequent renewal of cambium was observed as small segments instead of complete ring. They interconnect with existing cambium and form anastomosing network. Due to the renewal of small sectors, the secondary phloem formed by earlier cambial segments became enclosed within the secondary xylem and conjunctive tissue. Formation of vessels and sieve elements were found confined only to few fusiform cells while rest of the cambial sector exclusively produced conjunctive tissue on either side. The secondary xylem formed in the beginning of successive rings of xylem remained rayless while thick stems showed presence of heterocellular rays with several cells in height and width. Accumulation of starch along with the presence of nuclei in the xylem fibres even after deposition of the secondary wall is consistent feature in both the species.