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1 Gravity Resistance in Plants Graduate School of Science, Osaka City University Takayuki Hoson ABSTRACT Gravity resistance, a response that enables the plant to develop against the gravitational force, is a principal graviresponse in plants, comparable to gravitropism. However, only limited information has been obtained for this graviresponse. We have examined the nature and mechanism of gravity resistance by both ground-based and space experiments. The gravity signal may be perceived by mechanoreceptors (mechanosensitive ion channels) on the plasma membrane and it appears that amyloplast sedimentation in statocytes is not involved. Transformation and transduction of the perceived signal may be mediated by the structural or physiological continuum of microtubule cell membrane cell wall. The final step is to construct a tough body by increasing cell wall rigidity, which is brought about by modification of the metabolism of anti-gravitational wall polysaccharides and modification of the cell wall environment, especially ph. We need to clarify the details of gravity resistance by future space experiments. For this purpose, we should established the procedure of on-site analyses on orbit with new instruments and suitable plant materials such as mutants. 1 g Hoson 23, 26

2 gravity resistance 23, 25, Hoson and Soga 23, Hoson et al g STS nm

3 (1 3),(1 4) , UDPG Hoson 1998, 22

4 Masuda et al g 1 3, , Hoson 1998 ATP Ooume et al. 25

5 HPLC 1 g 199 g 1999, Hoson and Soga , , , , Hoson and Soga 23

6 1967 Biosatellite II STS-95 BRIC-RC RICE Hoson et al. 22, Soga et al , 22, Hoson et al Arabidopsis Rice Length (mm) Control Space Control Space Time (h)

7 STS-95 RICE Hoson et al. 22, Soga et al RICE 1 3, Hoson et al. 22, Soga et sl Arabidopsis 2. Rice Cell wall extensibility ( m/g) Control Space Control Space Time (h)

8 1 3, , 25, Hoson and Soga 23, Hoson et al , Hoson and Soga 23, Hoson et al. 25 g g 1 g g Length of hypocotyls (mm) R= Extensibility ( m/g) R = Log [g]

9 Elongation (mm) g 3 g 3 g 1 g Extensibility ( m/g) Time (h) , Hoson 1999 Kiss 2 Tasaka et al. 21 Boonsirichai et al. 22

10 Soga et al. 24 Soga et al. 25 Soga et al. 24, 25 25, Hoson and Soga 23, Hoson et al g

11 Wayne and Staves 1997 cdna -CoA HMGR - TUA3 Yoshioka et al. 23 HMGR HMGR Koizumi et al Matsumoto et al Soga et al. 26 HMGR Koizumi et al. 27, Matsumoto et al. 27 HMGR 1 g

12 Hypocotyl length (mm) g 3 g Diamter ( m) Wild hmg Wild hmg HMGR Hypocotyl length (mm) g 3 g Diamter ( m) Wild tua Wild tua -

13 1 g + H-ATPase 3 g ph.5.8 Soga et al. 2a 2b ph ph ph

14 ph ph Soga et al., 2a 2b Soga et al. 24, 25 MCA1 Nakagawa et al , Soga et al. 27

15 26 Nakabayashi et al g 1 g 1 g g 1 g

16 1 g 1 g 5% 1 g STS-95

17 Boonsirichai, K., Guan, C., Chen, R. and Masson, P. H. 22 Root gravitropism: an experimental tool to investigate basic cellular and molecular processes underlying mechanosensing and signal transmission in plants. Annu. Rev. Plant Biol. 53: Hoson, T Apoplast as the site of response to environmental signals. J. Plant Res. 111: Hoson, T Interaction of gravity with other environmental factors in growth and development: an introduction. Adv. Space Res. 23: Hoson,T.,Soga,K.,Mori,R.,Saiki,M.,Wakabayashi,K.,Kamisaka,S.,Kamigaichi,S.,Aizawa, S., Yoshizaki, I., Mukai, C., Shimazu, T., Fukui, K. and Yamashita, M Morphogenesis of rice and Arabidopsis seedlings in space. J. Plant Res. 112: : Hoson, T. 22 Physiological functions of plant cell coverings. J. Plant Res. 115: Hoson, T., Soga, K., Mori, R. Saiki, M., Nakamura, Y., Wakabayashi, K. and Kamisaka, S. 22 Stimulation of elongation growth and cell wall loosening in rice coleoptiles under microgravity conditions in space. Plant Cell Physiol. 43: : Hoson, T. 23 Development of the anti-gravitational system in land plants and its implication for the interaction between plants and other organisms. Biol. Sci. Space 17: Hoson, T. and Soga, K. 23 New aspects of gravity responses in plant cells. Int. Rev. Cytol. 229: : : Hoson, T., Saito, Y., Soga, K. and Wakabayashi, K. 25 Signal perception, transduction, and response in gravity resistance. Another graviresponse in plants. Adv. Space Res. 36: : Hoson, T. 26 The mechanism and significance of gravity resistance in plants. J. Gravit. Physiol. 13: Kiss, J. Z. 2 Mechanisms of the early phases of plant gravitropism. Critic. Rev. Plant Sci. 19:

18 Koizumi,T.,Sakaki,T.,Usui,S.,Soga,K.,Wakabayashi,K.andHoson,T. 27 Changesin membrane lipid composition in azuki bean epicotyls under hypergravity conditions: Possible role of membrane sterols in gravity resistance. Adv. Res. Space in press. MAPs. 2 p.3. Masuda, Y., Kamisaka, S. and Hoson, T Growth behaviour of rice coleoptiles. J. Plant Physiol. 152: Matsumoto, S., Saito, Y., Kumasaki, S., Soga, K., Wakabayashi, K. and Hoson, T. 27 Up-regulation of tubulin genes and roles of microtubules in hypergravity-induced growth modifications in Arabidopsis hypocotyls. Adv. Res. Space in press Nakabayashi, I., Karahara, I., Tamaoki, D., Masuda, K., Wakasugi, T., Yamada, K., Soga, K., Hoson, T. and Kamisaka, S. 26 Hypergravity stimulus enhances primary xylem development and decreases mechanical properties of secondary cell walls in inflorescence stems of Arabidopsis thaliana. Ann. Bot. 97: Nakagawa, Y., Katagiri, T., Shinozaki, K., Qi, Z., Tatsumi, H., Furuichi, T., Kishigami, A., Sokabe,M.,Kojima,I.,Sato,S.,Kato,T.,Tabata,S.,Iida,K.,Terashima,A.,Nakano,M., Ikeda, M., Yamanaka, T. and Iida, H. 27 Arabidopsis plasma membrane protein crucial 2+ for Ca influx and touch sensing in roots. Proc. Natl. Acad. Sci. USA in press p.31. Ooume, K., Soga, K., Wakabayashi, K. and Hoson, T. 25 Osmoregulation under condition of submergence in azuki bean epicotyls. Plant Cell Physiol. 46: s164. Soga, K., Wakabayashi, K., Hoson, T. and Kamisaka, S. 2a Changes in the apoplastic ph are involved in regulation of xyloglucan breakdown of azuki bean epicotyls under hypergravity conditions. Plant Cell Physiol. 41: Soga, K., Wakabayashi, K., Hoson, T. and Kamisaka, S. 2b Hypergravity-induced increase in the apoplastic ph and its possible involvement in suppression of -glucan breakdown in maize seedlings. Aust. J. Plant Physiol. 27: Soga, K., Wakabayashi, K., Kamisaka, S. and Hoson, T. 22 Stimulation of elongation growth and xyloglucan breakdown in Arabidopsis hypocotyls under microgravity conditions in space. Planta 215: Soga, K., Wakabayashi, K., Kamisaka, S. and Hoson, T. 24 Graviperception in growth inhibition of plant shoots under hypergravity conditions produced by centrifugation is independent of that in gravitropism and may involve mechanoreceptors. Planta 218:

19 Soga, K., Wakabayashi, K., Kamisaka, S. and Hoson, T. 25 Mechanoreceptors rather than sedimentable amyloplasts perceive the gravity signal in hypergravity-induced inhibition of root growth in azuki bean. Funct. Plant Biol. 32: Soga, K., Wakabayashi, K., Kamisaka, S. and Hoson, T. 26 Hypergravity induces reorientation of cortical microtubules and modifies growth anisotropy in azuki bean epicotyls. Planta 224: Soga, K. Arai, K., Wakabayashi, K., Kamisaka, S. and Hoson, T. 27 Modifications of xyloglucan metabolism in azuki bean epicotyls under hypergravity conditions. Adv. Res. Space in press Tasaka, M., Kato, T. and Fukaki, H. 21 Genetic regulation of gravitropism in higher plants. Int. Rev. Cytol. 26: Wayne, R. and Staves, M. P A down-to earth model of gravisensing. Gravi. Space Biol. Bull. 1: Yoshioka, R., Soga, K., Wakabayashi, K., Takeba, G. and Hoson, T. 23 Hypergravityinduced changes in gene expression in Arabidopsis hypocotyls. Adv. Space Res. 31:

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