The TRANSPARENT TESTA GLABRA1 ( TTG1 ) locus regulates several developmental and biochemical pathways in Arabidopsis, including the formation of hairs on leaves, stems, and roots, and the production of seed mucilage and anthocyanin pigments. The TTG1 locus has been isolated by positional cloning, and its identity was confirmed by complementation of a ttg1 mutant. The locus encodes a protein of 341 amino acid residues with four WD40 repeats. The protein is similar to AN11, a regulator of anthocyanin biosynthesis in petunia, and more distantly related to those of the ␤ subunits of heterotrimeric G proteins, which suggests a role for TTG1 in signal transduction to downstream transcription factors. The 1.5-kb TTG1 transcript is present in all major organs of Arabidopsis. Sequence analysis of six mutant alleles has identified base changes producing truncations or single amino acid changes in the TTG1 protein. INTRODUCTIONThe TRANSPARENT TESTA GLABRA1 ( TTG1 ) locus controls many apparently unrelated characters of Arabidopsis (catalogued by Koornneef, 1981), several of which appear to be confined to the epidermal cell layer of different tissues. ttg1 mutants have a glabrous phenotype, possessing none of the leaf or stem hairs (trichomes) that normally are derived from the meristematic L1 cell layer. Purple anthocyanin pigments are absent from the ttg1 seed coat, causing the transparent testa phenotype in which the yellow cotyledons are visible through the testa. In wild-type plants, anthocyanins are present in the hypocotyl of seedlings and in the stem and leaves of plants as they age, and they are inducible by many forms of stress, including high light, poor nutrients, or water stress. ttg1 mutants completely lack anthocyanins in the epidermis and in subepidermal layers of leaves and stems. Mucilage normally found in the cell wall of the seed coat is absent in ttg1 mutants. Seeds of ttg1 plants do not require drying and cold treatments to germinate and therefore exhibit an altered seed dormancy when compared with ecotypes, such as Landsberg erecta (L er ;Koornneef, 1981;Léon-Kloosterziel et al., 1994). This characteristic of ttg1 mutants may be linked to an altered seed coat structure. The TTG1 gene appears to have the opposite effect on root hair formation when compared with its effect on leaf hair initiation. In Arabidopsis, root hairs extend from root epidermal cells only in files of cells that contact two underlying cortical cells, whereas in ttg1 mutants, extra root hairs occur in the atrichoblast cell files (Galway et al., 1994). Under laboratory growth conditions, mutations at the ttg1 locus do not greatly affect the viability of the plants.In ttg1 mutants, the anthocyanin biosynthetic pathway is blocked at the dihydroflavonol-4-reductase (DFR) step, because DFR-encoding transcripts have not been detected in these mutants (Shirley et al., 1995). By contrast, transcripts of the chalcone synthase and chalcone isomerase genes are unaffected. The point of regulation of the pathway by TTG1 was confirmed by the clon...

The TRANSPARENT TESTA GLABRA1 (TTG1) locus regulates several developmental and biochemical pathways in Arabidopsis, including the formation of hairs on leaves, stems, and roots, and the production of seed mucilage and anthocyanin pigments. The TTG1 locus has been isolated by positional cloning, and its identity was confirmed by complementation of a ttg1 mutant. The locus encodes a protein of 341 amino acid residues with four WD40 repeats. The protein is similar to AN11, a regulator of anthocyanin biosynthesis in petunia, and more distantly related to those of the beta subunits of heterotrimeric G proteins, which suggests a role for TTG1 in signal transduction to downstream transcription factors. The 1.5-kb TTG1 transcript is present in all major organs of Arabidopsis. Sequence analysis of six mutant alleles has identified base changes producing truncations or single amino acid changes in the TTG1 protein.

Trichome development in Arabidopsis thaliana is a well-characterized model for the study of plant cell differentiation. Two genes that play an essential role in the initiation of trichome development are GL1 and TTG. Mutations in either gene prevent the initiation of most trichomes. The GL1 gene encodes a myb-related transcription factor. Mutations in TTG are pleiotropic, affecting anthocyanins, root hairs, and seed coat mucilage in addition to trichomes. Six ttg alleles were examined and shown to form a hypomorphic series. The severity of all aspects of the ttg phenotype varied in parallel in this allelic series. The weakest allele, ttg-10, causes frequent clusters of adjacent trichomes, suggesting a role for TTG in inhibiting neighboring cells from choosing the trichome fate. This allele results from a mutation in the 5′-untranslated region of ttg and creates an out-of-frame upstream AUG codon. The ttg-10 allele shows several unusual genetic interactions with the weak hypomorphic gl1-2 allele, including intergenic noncomplementation and a synthetic glabrous phenotype. These interactions are specific for the gl1-2 allele. The implication of these results for current models of trichome development is discussed.