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First published online January 28, 2005; 10.1104/pp.104.054494

Plant Physiology 137:663-670 (2005)
© 2005 American Society of Plant Biologists

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DEVELOPMENT AND HORMONE ACTION

Nitric Oxide Mediates Gravitropic Bending in Soybean Roots1,[w]

Xiangyang Hu2, Steven J. Neill, Zhangcheng Tang and Weiming Cai*

Institute of Plant Physiology and Ecology, Shanghai Institutes for Biological Sciences, Chinese Academy of Sciences, Graduate School of the Chinese Academy of Sciences, Shanghai 200032, China (X.H., Z.T., W.C.); and Centre for Research in Plant Science, University of the West of England, Bristol BS16 1QY, United Kingdom (S.J.N.)

Plant roots are gravitropic, detecting and responding to changes in orientation via differential growth that results in bending and reestablishment of downward growth. Recent data support the basics of the Cholodny-Went hypothesis, indicating that differential growth is due to redistribution of auxin to the lower sides of gravistimulated roots, but little is known regarding the molecular details of such effects. Here, we investigate auxin and gravity signal transduction by demonstrating that the endogenous signaling molecules nitric oxide (NO) and cGMP mediate responses to gravistimulation in primary roots of soybean (Glycine max). Horizontal orientation of soybean roots caused the accumulation of both NO and cGMP in the primary root tip. Fluorescence confocal microcopy revealed that the accumulation of NO was asymmetric, with NO concentrating in the lower side of the root. Removal of NO with an NO scavenger or inhibition of NO synthesis via NO synthase inhibitors or an inhibitor of nitrate reductase reduced both NO accumulation and gravitropic bending, indicating that NO synthesis was required for the gravitropic responses and that both NO synthase and nitrate reductase may contribute to the synthesis of the NO required. Auxin induced NO accumulation in root protoplasts and asymmetric NO accumulation in root tips. Gravistimulation, NO, and auxin also induced the accumulation of cGMP, a response inhibited by removal of NO or by inhibitors of guanylyl cyclase, compounds that also reduced gravitropic bending. Asymmetric NO accumulation and gravitropic bending were both inhibited by an auxin transport inhibitor, and the inhibition of bending was overcome by treatment with NO or 8-bromo-cGMP, a cell-permeable analog of cGMP. These data indicate that auxin-induced NO and cGMP mediate gravitropic curvature in soybean roots.


1 This work was supported by the Chinese Academy of Sciences (grant no. KSCX2–SW–322), the Shanghai Institute of Plant Physiology and Ecology, and the National Natural Sciences Foundation of China (grant no. 39770199). Collaboration between the United Kingdom and China was supported by The Royal Society and The Biotechnology and Biological Sciences Research Council.

2 Present address: Department of Botany and Plant Sciences, University of California, Riverside, CA 92521.

[w] The online version of this article contains Web-only data.

Article, publication date, and citation information can be found at www.plantphysiol.org/cgi/doi/10.1104/pp.104.054494.

* Corresponding author; e-mail wmcai{at}iris.sipp.ac.cn; fax 86–21–54924015.

Received October 8, 2004; returned for revision December 16, 2004; accepted December 20, 2004.


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