Aberrant cell expansion in the elongation mutants of barley

The elongation (elo) mutants of barley (Hordeum vulgare cv 'Himalaya') are a class of dwarf plants with defects affecting cell expansion. The phenotypes of mutants in three of the elo loci (elo1, elo2 and elo3) are recessive to the wild-type allele, and the mutations at elo-4 and elo-5 are semi-dominant. Allelism tests showed that elo1, elo2 and elo3 were at separate loci, and mapping data indicated that elo-5 was possibly allelic to either elo1 or elo2. A phenotype common to all elo mutants was the presence of short, radially swollen cells on the leaf epidermis, indicating a defect in longitudinal cell expansion. In three of the mutants, elo1, elo3 and elo5, this was accompanied by a twisting growth habit. Two of the mutations, elo2 and elo-5, affected cell division, with aberrant periclinal cell division resulting in the formation of increased cell layers in the leaf epidermis of elo2 and elo-5 homozygotes and in the aleurone layer of elo2 grains. Misplaced anticlinal divisions also occurred in the elo-5 leaf epidermis. Leaf cell walls of all elo lines contained less cellulose than the wild- type, and the cortical microtubules in elongating root epidermal cells in some elo lines were more randomly oriented than in the wild-type, consistent with the presence of radially swollen cells. We discuss possible functions for the Elo genes in primary cell wall synthesis.