Clonal analysis of the development of the barley (Hordeum vulgare L.) leaf using periclinal chlorophyll chimeras

Barley seedlings that show mosaic expression of chlorophyll were selected from the progenies of mutagenized seeds. The sectored plants were grown under conditions that lead to the formation of lateral tillers, and a fraction of these had different kinds of leaf variegation. These sectorially and periclinally chimeric shoots were used to analyze the cellular organization of the barley shoot apex and the clonal development of the leaf. The shoot apex is organized in two cell lineages, L1 and L2. As well as giving rise to the epidermis, the L1 layer contributes to leaf mesophyll, particularly at the margins, but, on the adaxial side of leaf laminae, also in more central positions. The L1 layer alone is responsible for the formation of the hood, a flower homologue structure present in strains homozygoous for the dominant allele at the It (hooded) locus. The relative contribution of L2 to leaf formation decreases in younger tillers and during tiller development from the basal to the flag leaf. Chimerism of the plants was generated by non-transmissible somatic events or by nuclear mutations.