The use of plants as integral components of life support systems remains a cornerstone of strategies for long-term human habitation of space and extraterrestrial colonization. Spaceflight experiments over the past few decades have refined the hardware required to grow plants in low-earth orbit and have illuminated fundamental issues regarding spaceflight effects on plant growth and development. Potential incipient hypoxia, resulting from the lack of convection-driven gas movement, has emerged as a possible major impact of microgravity. We developed transgenic Arabidopsis containing the alcohol dehydrogenase (Adh) gene promoter linked to the -glucuronidase (GUS) reporter gene to address specifically the possibility that spaceflight induces the plant hypoxia response and to assess whether any spaceflight response was similar to control terrestrial hypoxia-induced gene expression patterns. The staining patterns resulting from a 5-d mission on the orbiter Columbia during mission STS-93 indicate that the Adh/GUS reporter gene was activated in roots during the flight. However, the patterns of expression were not identical to terrestrial control inductions. Moreover, although terrestrial hypoxia induces Adh/GUS expression in the shoot apex, no apex staining was observed in the spaceflight plants. This indicates that either the normal hypoxia response signaling is impaired in spaceflight or that spaceflight inappropriately induces Adh/GUS activity for reasons other than hypoxia.
The principal objective of the research reported here was to determine whether a plant's periodic growth oscillations, called circumnutations, would persist in the absence of a significant gravitational or inertial force. The definitive experiment was made possible by access to the condition of protracted near weightlessness in an earth satellite. The experiment, performed during the first flight of Spacelab on the National Aeronautics and Space Administration shuttle, Columbia, in November and December, 1983, tested a biophysical model, proposed in 1967, that might account for circumnutation as a gravity-dependent growth response. However, circumnutations were observed in microgravity. They continued for many hours without stimulation by a significant g-force. Therefore, neither a gravitational nor an inertial g-force was an absolute requirement for initation or continuation of circumnutation. On average, circumnutation was significantly more vigorous in satellite orbit than on earth-based clinostats. Therefore, at least for sunflower (Helianthus annuus L.) circumnutation, clinostatting is not the functional equivalent of weightlessness.
For over half a century and especially since the 1960's a number of plant physiologists, seeking to explain the impressively ubiquitous mechanism that drives and regulates circumnutation in all growing plant organs, have been unable to agree on whether the differential growth process that leads to circumnutational oscillations is gravity dependent. There has been fairly general agreement that the question might be answered, if test plants could be deprived of all significant gravitational stimuli as would be possible in the near weightlessness or free fall environment of satellite orbit. Such an experiment was carried out during the Spacelab 1 mission. Circumnutational oscillations were observed which demonstrated that a protracted input of gravitational information from the environment was not required for initiation or maintenance of circumnutation in sunflower hypocotyls.
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