Developmental basis of homeosis in precociously germinating Brassica napus embryos: phase change at the shoot apex.

Precociously germinating Brassica napus (oilseed rape) embryos produce extra cotyledons or chimeric organs with sectors of cotyledon and leaf tissue, rather than leaves, at the shoot apex. To investigate this phenomenon in more detail, scanning electron microscopy was used to examine the development of organ primordia at the shoot apex. In situ hybridizations with molecular markers of the embryonic phase were used to assess the status of individual cells in the shoot apex with regard to the transition between embryonic and vegetative phases. The results indicate that, under conditions that support precocious germination, primordia develop at the shoot apex in the mode characteristic of postgerminative growth, i.e. they arise sequentially in a spiral phyllotaxy. Cells in the rest of the embryo, however, can continue to express molecular markers of the embryonic phase for several weeks after the start of culture. When patterns of gene expression and the fate of individual primordia were compared, a strong correlation was found between organ identity and the status of cells in the vicinity of the meristem with regard to phase. Primordia that develop in situations where neighboring cells are in the embryonic phase always produce organs with cotyledon morphology. Primordia that develop in situations where neighboring cells have exited the embryonic phase produce leaves. Based on an examination of situations where chimeric organs are produced, I propose that short range interactions or signalling are likely to be involved in communicating information about phase to developing primordia.