Homeodomain-leucine zipper (HD-Zip) proteins are putative transcription factors identified only in plants. Related Arabidopsis homeobox genes, isolated by virtue of sequence conservation within the helix-3 region of the homeodomain, fall into four families based on sequence similarity. This paper reports the characterization of Athb-10, a 747 amino acid protein belonging to the fourth HD-ZIP family. The studies indicate that, although less conserved, the leucine zipper of Athb-10 can functionally replace that of Athb-2 in an in vitro DNA-binding assay. Gene mapping experiments and sequence comparison analysis revealed that Athb-10 corresponds to GLABRA2, a homeodomain protein involved in trichome development. The mRNA expression analysis revealed that Athb-10/GLABRA2 is expressed not only in trichome-bearing organs, but also in the root. The analysis of wild-type and mutant plants showed that the Athb-10/GLABRA2 gene expression in the aerial part of the plant and in the root is affected by mutations at the TTG locus. Morphological analysis of the g/2-1 mutant revealed that the gene is necessary not only for local outgrowth of the trichome, but also for the regulation of root hair development in a subset of epidermal cells. Interestingly, the development of root hair cells in a position normally occupied by non-hair cells is dependent upon the ethylene regime in which the gl2-1 plants are grown. Sequence analysis of the gl2-1 allele revealed that the mutant gene encodes a truncated protein that might still retain a partial activity responsible for the formation of aborted trichomes and for the ethylene-dependent regulation of root hair formation.