Plant growth, development, and environmental responsiveness are dependent on hormone-induced gene expression. This dissertation reports multiple interactions between the plant hormones auxin and ethylene and investigates their contribution to the gravitropic response, elongation growth, adventitious root formation, callus and tracheary element initiation and growth, and flower development. Four mutants of tomato (Lycopersicon esculentum, Mill.) altered in either hormone production or hormone response were used to test the involvement of ethylene and auxin. These mutants included diageotropica (dgt) which is auxin-resistant, Never-ripe (Nr), which is ethylene-resistant, epinastic (epi), which overproduces ethylene and lazy-2 (lz-2), which exhibits a phytochrome-dependent reversed-gravitropic response. Additionally, a double mutant between Nr and dgt was constructed and tested. Gravitropism was studied as an exemplary process involving both auxin and ethylene. Mutant analysis demonstrated that ethylene does not play a primary role in the gravitropic response via the currently known ethylene response pathways. However, ethylene can modify the gravitropic response, e.g. the delayed gravitropic response of the dgt mutant can be restored with exceedingly low concentrations of ethylene and ethylene synthesis- and ethylene-action inhibitors can partially inhibit the graviresponse. The role of gravity in tracheary element (TE) production was tested in microgravity (during a space shuttle flight) and in hypergravity (centrifugation). A correlation was found between gravitational force and the production of TEs, with decreased numbers of TEs produced in microgravity and increased numbers produced in response to hypergravity. Increased production of TEs by dgt in both increased and reduced gravity indicates that gravity regulates vascular development via a DGT-dependent pathway involving auxin. Combination of both the Nr and dgt mutations in a double mutant leads to plants which exhibit the reduction of auxin-sensitivity typical of dgt as well as a delay in fruit ripening typical of Nr. The reduced gravitropic response of the dgt mutant was restored to wild-type levels in the double mutant confirming a complex role for ethylene in the gravitropic response. Abnormal floral organ development was observed in a subset of double mutant flowers.These data demonstrate multiple connections between auxin and ethylene during development and provide further insight into their cellular interactions. / Graduation date: 2001

The phytohormones auxin and cytokinin control plant development through a complex network of interactions which include synergistic, additive, and opposite effects whose mechanisms are unknown. The auxin-insensitive diageotropica (dgt) mutant provided a tool to dissect the relationship between auxin- and cytokinin-induced responses in tomato. Morphological, physiological, and molecular data support the proposal that auxin and cytokinins control a common set of developmental processes through separate signal transduction pathways which interact downstream from the DGT gene product. Morphological traits of dgt plants, such as reduced root and shoot growth, reduced leaf complexity, and reduced apical dominance were phenocopied by exogenous cytokinin application to wild-type plants, demonstrating that cytokinins and the DGT-mediated auxin response control a common set of phenotypic characteristics. Because the dgt mutation had no detectable effects on the levels of endogenous cytokinins or on the cytokinin sensitivity of whole plants, cytokinins were hypothesized to cause dgt-like effects on plant development through inhibiting auxin-induced responses. This hypothesis was supported by physiological experiments showing that auxin-induced elongation and ethylene synthesis were inhibited in cytokinin-treated wild-type and in untreated dgt hypocotyls. Differences between the effects of cytokinins and the dgt mutation on auxin responses became apparent at the molecular level. Experiments on the auxin-induced accumulation of transcripts for two ACC-synthase genes and one SAUR gene demonstrated that cytokinin treatment selectively reduced the auxin-induced expression of only one ACC-synthase gene, while the dgt mutation inhibited the auxin-inducibility of all three genes. The effects of the dgt mutation were thus more pleiotropic than the cytokinin effects, suggesting that cytokinins inhibit auxin-responses downstream from the DGT gene product. In vitro culture of dgt hypocotyl explants and calli demonstrated shared or interacting signal transduction pathways for auxin and cytokinin in the stimulation of cell division, and independent pathways for the control of organ regeneration and vascular differentiation. / Graduation date: 1997