How tightly land plants are adapted to the gravitational force (g) prevailing on Earth has been of interest because unlike many other environmental factors, g presents as a constant force. Ontogeny of mature angiosperms begins with an embryo that is formed after tip growth by a pollen tube delivers the sperm nucleus to the egg. Because of the importance to plant fitness, we have investigated how gravity affects these early stages of reproductive development. Arabidopsis thaliana (L.) Heynh. plants were grown for 13 days prior to being transferred to growth chambers attached to a large diameter rotor, where they were continuously exposed to 2-g or 4-g for the subsequent 11 days. Plants began flowering 1 day after start of the treatments, producing hundreds of flowers for analysis of reproductive development. At 4-g, Arabidopsis flowers self-pollinated normally but did not produce seeds, thus derailing the entire life cycle. Pollen viability and stigma esterase activity were not compromised by hypergravity; however, the growth of pollen tubes into the stigmas was curtailed at 4-g. In vitro pollen germination assays showed that 4-g average tube length was less than half that for 1-g controls. Closely related Brassica rapa L., which produces seeds at 4-g, required forces in excess of 6-g to slow in vitro tube growth to half that at 1-g. The results explain why seed production is absent in Arabidopsis at 4-g and point to species differences with regard to the g-sensitivity of pollen tube growth.
Previous experiments had shown that microgravity adversely affected seed development in Brassica rapa L. We tested the hypothesis that gravity controls seed development via modulation of gases around the developing seeds, by studying how hypergravity affects the silique microenvironment and seed development. Using an in vitro silique maturation system, we sampled internal silique gases for 16 d late in the seed maturation sequence at 4 g or 1 g. The carbon dioxide level was significantly higher inside the 4-g siliques, and the immature seeds became heavier than those maturing at 1 g. Pollination and early embryo development were also studied by growing whole plants at 2 g or 4 g for 16 d inside chambers mounted on a large-diameter centrifuge. Each day the rotor was briefly stopped to permit manual pollination of flowers, thereby producing cohorts of same-aged siliques for comparison with stationary control material. The loss of starch and soluble carbohydrates during seed development was accelerated in hypergravity, with seeds developing at 4 g more advanced by 2 d than those at 1 g. Seeds produced at 4 g contained more lipid than those at 1 g. Taken together, these results indicate that hypergravity enhances gas availability to the developing embryos. Gravity's role in seed development is of importance to the space programme because of the plan to use plants for food production and habitat regeneration in extraterrestrial settings. These results are significant because they underscore the tight co-regulation of Brassica seed development and the atmosphere maintained inside the siliques.
How gravity influences the growth form and flavor components of plants is of interest to the space program because plants could be used for food and life support during prolonged missions away from the planet, where that constant feature of Earth's environment does not prevail. We used plant growth hardware from prior experiments on the space shuttle to grow Brassica rapa and Arabidopsis thaliana plants during 16-d or 11-d hypergravity treatments on large-diameter centrifuge rotors. Both species showed radical changes in growth form, becoming more prostrate with increasing g-loads (2-g and 4-g). In Brassica, height decreased and stems thickened in a linear relationship with increasing g-load. Glucosinolates, secondary compounds that contribute flavor to Brassica, decreased by 140% over the range of micro to 4-g, while the structural secondary compound, lignin, remained constant at ∼15% (w/w) cell wall dry mass. Stem thickening at 4-g was associated with substantial increases in cell size (47%, 226%, and 33% for pith, cortex, and vascular tissue), rather than any change in cell number. The results, which demonstrate the profound effect of gravity on plant growth form and secondary metabolism, are discussed in the context of similar thigmostresses such as touch and wind.
The extent of seed transmission of dry-rot was assessed over a period of three years by sowing seed lines with known levels of infection and noting the amount of dry-rot which subsequently developed in each line.
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