The control of cell fate was investigated in the root epidermis of Arabidopsis thaliana. Two distinct types of differentiated epidermal cells are normally present: root-hair-bearing cells and hairless cells. In wild-type Arabidopsis roots, epidermal cell fate was found to be correlated with cell position, with root-hair cells located over radial walls between cortical cells, and with hairless cells located directly over cortical cells. This normal positional relationship was absent in ttg (transparent testa glabrous) mutants (lacking trichomes, anthocyanins, and seed coat mucilage); epidermal cells in all positions differentiate into root-hair cells. The opposite condition was generated in roots of transgenic Arabidopsis expressing the maize R (R-Lc) gene product (a putative TTG homologue) under the control of a strong promoter (CaMV35S), which produced hairless epidermal cells in all positions. In both the ttg and R-expressing roots, epidermal cell differentiation was affected at an early stage, prior to the onset of cell elongation or root-hair formation. The ttg mutations were also associated with abnormalities in the morphology and organization of cells within and surrounding the root apical meristem. The results indicate that alterations in TTG activity cause developing epidermal cells to misinterpret their position and differentiate into inappropriate cell types. This suggests that, in wild-type roots, TTG provides, or responds to, positional signals to cause differentiating epidermal cells that lie over cortical cells to adopt a hairless cell fate.

Root-hair initiation inArabidopsis thaliana provides a model for studying cell polarity and its role in plant morphogenesis. Root hairs normally emerge at the apical end of root epidermal cells, implying that these cells are polarized. We have identified a mutant, rhd6, that displays three defects: (a) a reduction in the number of root hairs, (b) an overall basal shift in the site of roothair emergence, and (c) a relatively high frequency of epidermal cells with multiple root hairs. These defects implicate the RHD6 gene in root-hair initiation and indicate that RHD6 is normally associated with the establishment of, or response to, root epidermal cell polarity. Similar alterations in the site of root-hair emergence, although less extreme, were also discovered in roots of the auxin-, ethylene-, abscisic acid-resistant mutant axr2 and the ethylene-resistant mutant etrl. All three rhd6 mutant phenotypes were rescued when either auxin (indoleacetic acid) or an ethylene precursor (1 -aminocyclopropane-1-carboxylic acid) was included in the growth medium. The rhd6 root phenotypes could be phenocopied by treating wild-type seedlings with an inhibitor of the ethylene pathway (aminoethoxyvinylglycine). These results indicate that RHD6 is normally involved in directing the selection or assembly of the root-hair initiation site through a process involving auxin and ethylene.

The Arabidopsis root produces a position-dependent pattern of hair-bearing and hairless cell types during epidermis development. have been reported to affect the production of root hair and hairless cells in the Arabidopsis root. In this study, genetic, molecular, and physiological tests were employed to define the roles of these loci and hormones. Epistasis tests and reporter gene studies indicated that the hairless cell-promoting genes TTG and GL2 are likely to act early to negatively regulate the ethylene and auxin pathways. Studies of the developmental timing of the hormone effects indicated that ethylene and auxin pathways promote root hair outgrowth after cell-type differentiation has been initiated. The genetic analysis of ethylene-and auxin-related mutations showed that root hair formation is influenced by a network of hormone pathways, including a partially redundant ethylene signaling pathway. A model is proposed in which the patterning of root epidermal cells in Arabidopsis is regulated by the cell position-dependent action of the TTGlGL2 pathway, and the ethylene and auxin hormone pathways act to promote root hair outgrowth at a relatively late stage of differentiation.

The Arabidopsis root produces a position-dependent pattern of hair-bearing and hairless cell types during epidermis development. Five loci (TRANSPARENT TESTA GLABRA [TTG], GLABRA2 [GL2], ROOT HAIR DEFECTIVE6 [RHD6], CONSTITUTIVE TRIPLE RESPONSE1 [CTR1], and AUXIN RESISTANT2 [AXR2]) and the plant hormones ethylene and auxin have been reported to affect the production of root hair and hairless cells in the Arabidopsis root. In this study, genetic, molecular, and physiological tests were employed to define the roles of these loci and hormones. Epistasis tests and reporter gene studies indicated that the hairless cell-promoting genes TTG and GL2 are likely to act early to negatively regulate the ethylene and auxin pathways. Studies of the developmental timing of the hormone effects indicated that ethylene and auxin pathways promote root hair outgrowth after cell-type differentiation has been initiated. The genetic analysis of ethylene-and auxin-related mutations showed that root hair formation is influenced by a network of hormone pathways, including a partially redundant ethylene signaling pathway. A model is proposed in which the patterning of root epidermal cells in Arabidopsis is regulated by the cell position-dependent action of the TTG/GL2 pathway, and the ethylene and auxin hormone pathways act to promote root hair outgrowth at a relatively late stage of differentiation.

The pattern of structures on most of the adult cuticle of Drosophila is determined in the larval imaginal disks. The Drosophila growth factor homolog decapentaplegic [dpp] is believed to participate in pattern formation in imaginal disks, primarily along what will become the proximal-to-distal axis of adult appendages. We report that dpp expression in wing, leg, and eye-antennal imaginal disks is localized to a band of cells along the presumptive proximal-to-distal axis. The pattern and level of dpp expression in imaginal disks is affected by mutant lesions that remove 3' cis-regulatory sequences. We demonstrate that one portion of the 3' cisregulatory region contains regulatory elements sufficient to activate gene expression in a subset of the cells that normally express dpp in the imaginal disks, allowing rescue of dpp mutant phenotypes. We propose that the complete dpp expression pattern is generated by an array of 3' regulatory elements that differ in their potency in specific disks and in certain positions within a disk. The identification of the factors that activate these elements should provide clues as to how positional information is encoded in imaginal disks.

The Organization for Economic Co-operation and Development (OECD) recommends the measurement of specific plant components for compositional assessments of new biotechnology-derived crops. These components include proximates, nutrients, antinutrients, and certain crop-specific secondary metabolites. A considerable literature on the natural variability of these components in conventional and biotechnology-derived crops now exists. Yet the OECD consensus also suggests measurements of any metabolites that may be directly associated with a newly introduced trait. Therefore, steps have been initiated to assess natural variation in metabolites not typically included in the OECD consensus but which might reasonably be expected to be affected by new traits addressing, for example, nutritional enhancement or improved stress tolerance. The compositional study reported here extended across a diverse genetic range of maize hybrids derived from 48 inbreds crossed against two different testers. These were grown at three different, but geographically similar, locations in the United States. In addition to OECD analytes such as proximates, total amino acids and free fatty acids, the levels of free amino acids, sugars, organic acids, and selected stress metabolites in harvested grain were assessed. The major free amino acids identified were asparagine, aspartate, glutamate, and proline. The major sugars were sucrose, glucose, and fructose. The most predominant organic acid was citric acid, with only minor amounts of other organic acids detected. The impact of genetic background and location was assessed for all components. Overall, natural variation in free amino acids, sugars, and organic acids appeared to be markedly higher than that observed for the OECD analytes.

l h e expansion of both root hairs and pollen tubes occurs by a process known as tip growth. In this report, an Arabidopsis thaliana mutant ( t i p l ) is described that displays defects in both root-hair and pollen-tube growth. l h e root hairs of the fipl mutant plants are shorter than those of the wild-type plants and branched at their base. l h e tipl pollen-tube growth defect was identified by the aberrant segregation ratio of phenotypically normal to mutant seeds in siliques from self-pollinated, heterozygous plants. Homozygous mutant seeds are not randomly distributed in the siliques, comprising only 14.4% of the total seeds, 5.3% of the seeds from the bottom half, and 2.2% of the seeds from the bottom quarter of the heterozygous siliques. Studies of pollen-tube growth in vivo showed that mutant pollen tubes grow more slowly than wild-type pollen through the transmitting tissue of wild-type flowers. Cosegregation studies indicate that the root-hair and pollen-tube defects are caused by the same genetic lesion. Based on these findings, the TlPl gene is likely to encode a product involved in a fundamental aspect of tip growth in plant cells.

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