[Changes in the mechanical and chemical properties of rice coleoptile cell walls under microgravity conditions].
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Biomedical subjects
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The effects of simulated microgravity conditions produced by a horizontal clinostat on the entire life cycle of Arabidopsis thaliana ecotype Columbia and Landsberg erecta were studied. Horizontal clinorotation affected little germination of seeds, growth and development of rosette leaves and roots during early vegetative growth stage, and the onset of the bolting of inflorescence axis and flower formation in reproductive growth stage, although it suppressed elongation of inflorescence axes. The clinorotation substantially reduced the numbers of siliques and seeds in Landsberg erecta, and completely inhibited seed production in Columbia. Seeds produced in Landsberg erecta on the clinostat were capable of germinating and developing rosette leaves normally on the ground. On the other hand, growth of pin formed mutant (pin/pin) of Arabidopsis ecotype Enkheim, which has a unique structure of inflorescence axis with no flower and extremely low levels of auxin polar transport activity, was inhibited and the seedlings frequently died during vegetative stage on the clinostat. Seed formation and inflorescence growth of the seedlings with normal shape (pin/+ or +/+) were also suppressed on the clinostat. These results suggest that the growth and development of Arabidopsis, especially in reproductive growth stage, is suppressed under simulated microgravity conditions on a clinostat. To complete the life cycle probably seems to be quite difficult, although it is possible in some ecotypes.
Elongation growth of dark grown maize (Zea mays L cv. Cross Bantam T51) coleoptiles and mesocotyls was suppressed by hypergravity at 30 g and above. Acceleration at 300 g significantly decreased the mechanical extensibility of cell walls of both organs. Hypergravity increased the amounts of hemicellulose and cellulose per unit length in mesocotyl walls, but not in coleoptile walls. The weight average molecular masses of hemicellulosic polysaccharides were also increased by hypergravity in both organs. On the other hand, the activities of beta-glucanases extracted from coleoptile and mesocotyl cell walls were decreased by hypergravity. These results suggest that the decreased activities of beta-glucanases by hypergravity cause an increase in the molecular mass of hemicellulosic polysaccharides of both organs. The upshift of molecular mass of hemicellulosic polysaccharides as well as the thickening of cell walls under hypergravity conditions seems to be involved in making the cell wall mechanically rigid, thereby inhibiting elongation growth of maize coleoptiles and mesocotyls.
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Seedlings of rice (Oryza saliva L.) and Arabidopsis (A. thaliana L.) were cultivated for 68.5 hr in the RICE experiment on board during Space Shuttle STS 95 mission, and changes in their growth and morphology were analyzed. Microgravity in space stimulated elongation growth of both rice coleoptiles and Arabidopsis hypocotyls by making their cell walls extensible. In space, rice coleoptiles showed an inclination toward the caryopsis in the basal region and also a spontaneous curvature in the same direction in the elongating region. These inclinations and curvatures were more prominent in the Koshihikari cultivar compared to a dwarf cultivar, Tan-ginbozu. Rice roots elongated in various directions including into the air on orbit, but two thirds of the roots formed a constant angle with the axis of the caryopsis. In space, Arabidopsis hypocotyls also elongated in a variety of directions and about 10% of the hypocotyls grew into the agar medium. No clear curvatures were observed in the elongating region of Arabidopsis hypocotyls. Such a morphology of both types of seedlings was fundamentally similar to that observed on a 3 D clinostat. Thus, it was confirmed by the RICE experiment that rice and Arabidopsis seedlings perform an automorphogenesis under not only simulated but also true microgravity conditions.
Photoperiodically controlled adult diapause in the bean bug, Riptortus clavatus (Thunberg) (Heteroptera: Alydidae) was due to suppression of corpus allatum activity under short-day conditions. The mechanical extensibility of the cuticle of the pronotum was significantly higher in nondiapause adults reared under long-day conditions than in diapause adults reared under short-day conditions. Furthermore, diapause adults accumulated significantly larger amount of lipids than nondiapause ones. It was then examined whether these two characteristics of adult diapause also depend on activity of the corpus allatum, by removal of the corpus allatum and transection of the nervi corporis allati. Even after these two kinds of surgery, adults responded to photoperiod and showed similar differences both in mechanical properties of the cuticle and in lipid content between long-day and short-day conditions. Therefore, inactivity of the corpus allatum is not responsible for the stiffer cuticle or higher lipid accumulation in diapause adults.
The life of plants and other organisms is governed by the constant force of gravity on earth. The mechanism of graviperception, signal transduction, and gravireaction is one of the major themes in space biology. When gravity controls each step of the life cycle such as growth and development, it does not work alone but operates with the interaction of other environmental factors. In order to understand the role of gravity in regulation of the life cycle, such interactions also should be clarified. Under microgravity conditions in space, various changes are brought about in the process of growth and development. Some changes would be advantageous to organisms, but others would be unfavorable. For overcoming such disadvantages, it may be required to exploit some other environmental factors which substitute for gravity in some properties. In terrestrial plants, gravity can be replaced by light under certain conditions. The gravity-substituting factors may play a principal role in future space development.
Activity of auxin polar transport in inflorescence axes of Arabidopsis thaliana grown under simulated microgravity conditions was studied in relation to the growth and development. Seeds were germinated and allowed to grow on an agar medium in test tubes on a horizontal clinostat. Horizontal clinostat rotation substantially reduced the growth of inflorescence axes and the productivity of seeds of Arabidopsis thaliana (ecotypes Landsberg erecta and Columbia), although it little affected seed germination, development of rosette leaves and flowering. The activity of auxin polar transport in inflorescence axes decreased when Arabidopsis plants were grown on a horizontal clinostat from germination stage, being ca. 60% of 1 g control. On the other hand, the auxin polar transport in inflorescence axes of Arabidopsis grown in 1 g conditions was not affected when the segments were exposed to various gravistimuli, including 3-dimensional clinorotation, during transport experiments. Pin-formed mutant of Arabidopsis, having a unique structure of the inflorescence axis with no flower and extremely low levels of the activity of auxin polar transport in inflorescence axes and endogenous auxin, did not continue its vegetative growth under clinostat rotation. These facts suggest that the development of the system of auxin polar transport in Arabidopsis is affected by microgravity, resulting in the inhibition of growth and development, especially during reproductive growth.
Airtight vessels have various advantages for space experiments. However, Arabidopsis thaliana plants scarcely produced seeds when grown in such vessels. The mechanism by which reproductive growth is inhibited in airtight vessels was studied. The length of the flower stalk was shorter when the plants were grown in airtight vessels. Thus, there was a possibility that the inhibition of reproductive growth was due to the inhibition of vegetative growth. However, even when the plants which has grown under non-airtight conditions and has reached to the flowering stage were transferred to airtight vessels, silique formation was inhibited, suggesting that the airtight environment directly influences reproductive growth. In airtight vessels, anther dehiscence was inhibited, which appears to be the cause of inhibition of silique formation and seed development. Reproductive growth recovered when silica gel was added to the vessels. These results suggest that in airtight vessels, high humidity causes a suppression of anther dehiscence, resulting in the inhibition of reproductive growth. Therefore, the control of humidity by ventilation should be taken into consideration in designing a growth chamber for space experiments.
Elongation growth of dark-grown azuki bean (Vigna angularis Ohwi et Ohashi cv. Takara) epicotyls was suppressed by hypergravity at 30 x g and above. Acceleration at 300 x g significantly decreased the mechanical extensibility of cell walls. The amounts of cell wall polysaccharides (pectin, hemicellulose-II and cellulose) per unit length of epicotyls increased under the hypergravity condition. Hypergravity also increased the amounts and the weight-average molecular mass of xyloglucans in the hemicellulose-II fraction, while decreasing the activity of xyloglucan-degrading enzymes extracted from epicotyl cell walls. These results suggest that hypergravity increases the amounts and the molecular mass of xyloglucans by decreasing xyloglucan-degrading activity. Modification of xyloglucan metabolism as well as the thickening of cell walls under hypergravity conditions seems to be involved in making the cell wall mechanically rigid, thereby inhibiting elongation growth of azuki bean epicotyls.
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