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        hd1, a Major Photoperiod Sensitivity Quantitative Trait Locus in Rice, Is Closely Related to the Ara

        The Plant Cell, Vol. 12, 2473–2483, December 2000, http://www.capitolkaraoke.com/doc/6badba0f6c85ec3a87c2c59b.html ? 2000 American Society of Plant Physiologists

        Hd1, a Major Photoperiod Sensitivity Quantitative Trait Locus in Rice, Is Closely Related to the Arabidopsis Flowering Time Gene CONSTANS

        Masahiro Yano,a,1,2 Yuichi Katayose,a,1 Motoyuki Ashikari,b Utako Yamanouchi,c,d Lisa Monna,c

        Takuichi Fuse,b Tomoya Baba,c Kimiko Yamamoto,c Yosuke Umehara,a Yoshiaki Nagamura,a

        and Takuji Sasaki a

        a National Institute of Agrobiological Resources, Tsukuba, Ibaraki 305-8602, Japan

        b Bio-oriented Technology Research Advancement Institution, Omiya, Saitama 331-8537, Japan

        c Institute of the Society for Techno-innovation of Agriculture, Forestry an

        d Fisheries, Tsukuba, Ibaraki 305-0854, Japan

        d Department of Biology, Faculty of Science, Toyama University, Toyama, 930-8555, Japan

        A major quantitative trait locus (QTL) controlling response to photoperiod, Hd1, was identified by means of a map-based cloning strategy. High-resolution mapping using 1505 segregants enabled us to define a genomic region of ?12 kb as a candidate for Hd1. Further analysis revealed that the Hd1 QTL corresponds to a gene that is a homolog of CON-STANS in Arabidopsis. Sequencing analysis revealed a 43-bp deletion in the first exon of the photoperiod sensitivity 1 (se1) mutant HS66 and a 433-bp insertion in the intron in mutant HS110. Se1 is allelic to the Hd1 QTL, as determined by analysis of two se1 mutants, HS66 and HS110. Genetic complementation analysis proved the function of the candidate gene. The amount of Hd1 mRNA was not greatly affected by a change in length of the photoperiod. We suggest that Hd1 functions in the promotion of heading under short-day conditions and in inhibition under long-day conditions.


        The transition of the apical meristem from vegetative to re-productive growth is a critical event in the life cycle of a plant. In rice, the timing of this transition affects the timing of heading. This timing, or heading date, is one of the critical traits considered for adapting rice to different cultivation ar-eas and cropping seasons. Rice is a short-day (SD) plant; its heading is promoted by short photoperiods. The response of the plant to length of day (referred to as photoperiod sen-sitivity [PS]) and its basic vegetative growth determine the heading date of rice. Many genetic studies of heading date have been performed, and several genes controlling PS in rice have been genetically identified, including Se1(Lm), Se3 to Se7, and E1 to E3 (Yokoo et al., 1980; Yamagata et al., 1986; Poonyarit et al., 1989; Sano, 1992; Yokoo and Okuno, 1993; Tsai, 1995; Kinoshita, 1998). H owever, only one gene involving photoperiod response in rice has been cloned, Se5 (Izawa et al., 2000). Little is known about the structure and function of PS genes in rice at the molecular level. In contrast, several genes involved in flowering time have been isolated, allowing clarification of part of the ge-netic control pathway in Arabidopsis (reviewed by Levy and Dean, 1998; Fowler et al., 1999; Kobayashi et al., 1999; Sheldon et al., 1999). Identification of the genes involved in flowering time has made it possible to determine the genetic control pathways for the response to photoperiod and ver-nalization in Arabidopsis (reviewed by Levy and Dean, 1998; Samach and Coupland, 2000). In addition, homologs of Ara-bidopsis genes for flowering time also function in Brassica napus (Robert et al., 1998).

        The major genes or quantitative trait loci (QTLs) for head-ing date have been mapped by using DNA markers in rice (Mackill et al., 1993; Li et al., 1995; Xiao et al., 1996; Yano et al., 1997; Lin et al., 1998; Tamura et al., 1998). Four QTLs for heading date involved in PS were mapped precisely as sin-gle Mendelian factors by using advanced backcross prog-eny (Yamamoto et al., 1998, 2000; Lin et al., 2000). A major gene controlling PS, Se1, first was identified as a naturally occurring variant (Yokoo et al., 1980) and was thought to lie at the same locus as the PS QTL Hd1, based on a compari-son of their map locations (Yano et al., 1997; Tamura et al., 1998; Yamamoto et al., 1998). Mutant lines with less PS were induced, and some were caused by mutations in the Se1 locus (Inoue et al., 1992). However, no direct evidence was available to prove their allelic relationship, and isolation

        1These authors contributed equally to this work.

        2To whom correspondence should be addressed. E-mail myano@ abr.affrc.go.jp; fax 81-298-38-7468.