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.
Chemical gradients and structural features within the pistil have been previously proposed as factors determining the directionality of pollen tube growth. In this study, we examine the behavior of pollen of eight species germinated in a dynamic oxygen gradient. While the germination rates of some species decreased directly with decreasing oxygen tension, other species showed no decrease in germination at oxygen tensions as low as 2 kPa. In one species, germination was consistently greater at decreased oxygen tensions than at ambient atmospheric levels. In three of the eight species tested, the developing pollen tube showed clear directional growth away from the more-oxygenated regions of the growth medium, while in one species growth was towards the more-oxygenated region. The remaining four species showed random tube growth. The pattern of oxytropic responses among the taxa suggests that this tropic behavior is both widespread and phylogenetically unpredictable.
Seeds developing within a locular space inside hollow fruit experience chronic exposure to a unique gaseous environment. Using two pepper cultivars, `Triton' (sweet) and `PI 140367' (hot), we investigated how the development of seeds is affected by the gases surrounding them. The atmospheric composition of the seed environment was characterized during development by analysis of samples withdrawn from the fruit locule with a gas-tight syringe. As seed weight plateaued during development, the seed environment reached its lowest O2 concentration (19%) and highest CO2 concentration (3%). We experimentally manipulated the seed environment by passing different humidified gas mixtures through the fruit locule at a rate of 60 to 90 mL·min-1. A synthetic atmosphere containing 3% CO2, 21% O2, and 76% N2 was used to represent a standard seed environment. Seeds developing inside locules supplied with this mixture had enhanced average seed weight, characterized by lower variation than in the no-flow controls due to fewer low-weight seeds. The importance of O2 in the seed microenvironment was demonstrated by reduction in seed weight when the synthetic atmosphere contained only 15% O2 and by complete arrest of embryo development when O2 was omitted from the seed atmosphere. Removal of CO2 from the synthetic atmosphere had no effect on seed weight, however, the CO2-free treatment accelerated fruit ripening by 4 days in the hot pepper. In the sweet peppers, fruit wall starch and sucrose were reduced by the CO2-free treatment. The results demonstrate that accretionary seed growth is being limited in pepper by O2 availability and suggest that variation in seed quality is attributable to localized limitations in O2 supply.
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