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ABSTRACT Plants must coordinate the regulation of biochemistry and anatomy to optimize photosynthesis and water-use efficiency. The formation of stomata, epidermal pores that facilitate gas
exchange, is highly coordinated with other aspects of photosynthetic development. The signalling pathways controlling stomata development are not fully understood1,2, although
mitogen-activated protein kinase (MAPK) signalling is known to have key roles. Here we demonstrate in _Arabidopsis_ that brassinosteroid regulates stomatal development by activating the MAPK
kinase kinase (MAPKKK) YDA (also known as YODA). Genetic analyses indicate that receptor kinase-mediated brassinosteroid signalling inhibits stomatal development through the glycogen
synthase kinase 3 (GSK3)-like kinase BIN2, and BIN2 acts upstream of YDA but downstream of the ERECTA family of receptor kinases. Complementary _in vitro_ and _in vivo_ assays show that BIN2
phosphorylates YDA to inhibit YDA phosphorylation of its substrate MKK4, and that activities of downstream MAPKs are reduced in brassinosteroid-deficient mutants but increased by treatment
with either brassinosteroid or GSK3-kinase inhibitor. Our results indicate that brassinosteroid inhibits stomatal development by alleviating GSK3-mediated inhibition of this MAPK module,
providing two key links; that of a plant MAPKKK to its upstream regulators and of brassinosteroid to a specific developmental output. Access through your institution Buy or subscribe This is
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Open access 04 May 2022 CHANGE HISTORY * _ 08 FEBRUARY 2012 The original supplementary information file posted online was incorrect and has been replaced. _ REFERENCES * Bergmann, D. C.
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proteomics. _Plant J._ 45, 616–629 (2006) Article CAS Google Scholar Download references ACKNOWLEDGEMENTS We thank K. Torii for providing seeds of the _er erl1 erl2_ triple mutant and
_scrm-D_ mutant, J.-M. Zhou for providing the _HOPAI1_ transgenic line and J. Li for providing the _bin2-3 bil1 bil2_ triple mutant. Research was primarily supported by a grant from the
National Institutes of Health (R01GM066258), and partially supported by the US Department of Energy (DE-FG02-08ER15973) and the Herman Frasch Foundation. D.C.B. is an investigator of the
Howard Hughes Medical Institute. AUTHOR INFORMATION AUTHORS AND AFFILIATIONS * Department of Plant Biology, Carnegie Institution for Science, Stanford, 94305-4150, California, USA Tae-Wuk
Kim & Zhi-Yong Wang * Department of Life Science, Hanyang University, Seoul 133-791, South Korea, Tae-Wuk Kim * Department of Biology, Stanford University, Stanford, 94305-5020,
California, USA Marta Michniewicz & Dominique C. Bergmann Authors * Tae-Wuk Kim View author publications You can also search for this author inPubMed Google Scholar * Marta Michniewicz
View author publications You can also search for this author inPubMed Google Scholar * Dominique C. Bergmann View author publications You can also search for this author inPubMed Google
Scholar * Zhi-Yong Wang View author publications You can also search for this author inPubMed Google Scholar CONTRIBUTIONS T.-W.K. performed all experiments. T.-W.K. and Z.-Y.W. designed the
experiments, analysed data and wrote the manuscript. M.M. cloned complementary DNAs of BSL2 and BSL3, and BSL2pro-BSL2. D.C.B. contributed materials and wrote the manuscript. CORRESPONDING
AUTHOR Correspondence to Zhi-Yong Wang. ETHICS DECLARATIONS COMPETING INTERESTS The authors declare no competing financial interests. SUPPLEMENTARY INFORMATION SUPPLEMENTARY INFORMATION This
file contains Supplementary Figures 1-13 with legends. _The original file posted online was incorrect and was replaced on 8 February 2012._ (PDF 4371 kb) POWERPOINT SLIDES POWERPOINT SLIDE
FOR FIG. 1 POWERPOINT SLIDE FOR FIG. 2 POWERPOINT SLIDE FOR FIG. 3 RIGHTS AND PERMISSIONS Reprints and permissions ABOUT THIS ARTICLE CITE THIS ARTICLE Kim, TW., Michniewicz, M., Bergmann,
D. _et al._ Brassinosteroid regulates stomatal development by GSK3-mediated inhibition of a MAPK pathway. _Nature_ 482, 419–422 (2012). https://doi.org/10.1038/nature10794 Download citation
* Received: 18 March 2011 * Accepted: 13 December 2011 * Published: 05 February 2012 * Issue Date: 16 February 2012 * DOI: https://doi.org/10.1038/nature10794 SHARE THIS ARTICLE Anyone you
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