First published online July 8, 2005; 10.1105/tpc.105.033464
The Plant Cell 17:2255-2270 (2005)
© 2005 American Society of Plant Biologists
Distinct Roles of GIGANTEA in Promoting Flowering and Regulating Circadian Rhythms in Arabidopsis
Tsuyoshi Mizoguchia,1,
Louisa Wrightb,1,
Sumire Fujiwaraa,1,
Frédéric Cremerb,
Karen Leec,
Hitoshi Onouchic,
Aidyn Mouradovb,
Sarah Fowlerd,
Hiroshi Kamadaa,
Joanna Putterilld and
George Couplandb,2
a Institute of Biological Sciences, University of Tsukuba, Tsukuba, Ibaraki 305-8572, Japan
b Max Planck Institute for Plant Breeding, D-50829 Cologne, Germany
c John Innes Centre, Norwich, United Kingdom
d School of Biological Sciences, University of Auckland, Private Bag 92019, Auckland, New Zealand
2 To whom correspondence should be addressed. E-mail coupland{at}mpiz-koeln.mpg.de; fax 49-221-5062-207.
The circadian clock acts as the timekeeping mechanism in photoperiodism. In Arabidopsis thaliana, a circadian clockcontrolled flowering pathway comprising the genes GIGANTEA (GI), CONSTANS (CO), and FLOWERING LOCUS T (FT) promotes flowering specifically under long days. Within this pathway, GI regulates circadian rhythms and flowering and acts earlier in the hierarchy than CO and FT, suggesting that GI might regulate flowering indirectly by affecting the control of circadian rhythms. We studied the relationship between the roles of GI in flowering and the circadian clock using late elongated hypocotyl circadian clock associated1 double mutants, which are impaired in circadian clock function, plants overexpressing GI (35S:GI), and gi mutants. These experiments demonstrated that GI acts between the circadian oscillator and CO to promote flowering by increasing CO and FT mRNA abundance. In addition, circadian rhythms in expression of genes that do not control flowering are altered in 35S:GI and gi mutant plants under continuous light and continuous darkness, and the phase of expression of these genes is changed under diurnal cycles. Therefore, GI plays a general role in controlling circadian rhythms, and this is different from its effect on the amplitude of expression of CO and FT. Functional GI:green fluorescent protein is localized to the nucleus in transgenic Arabidopsis plants, supporting the idea that GI regulates flowering in the nucleus. We propose that the effect of GI on flowering is not an indirect effect of its role in circadian clock regulation, but rather that GI also acts in the nucleus to more directly promote the expression of flowering-time genes.
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