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      A tomato HD-Zip homeobox protein, LeHB-1, plays an important role in floral organogenesis and ripening

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          Abstract

          Ethylene is required for climacteric fruit ripening. Inhibition of ethylene biosynthesis genes, 1-aminocyclopropane-1-carboxylate (ACC) synthase and ACC oxidase, prevents or delays ripening, but it is not known how these genes are modulated during normal development. LeHB-1, a previously uncharacterized tomato homeobox protein, was shown by gel retardation assay to interact with the promoter of LeACO1, an ACC oxidase gene expressed during ripening. Inhibition of LeHB-1 mRNA accumulation in tomato fruit, using virus-induced gene silencing, greatly reduced LeACO1 mRNA levels, and inhibited ripening. Conversely, ectopic overexpression of LeHB-1 by viral delivery to developing flowers elsewhere on injected plants triggered altered floral organ morphology, including production of multiple flowers within one sepal whorl, fusion of sepals and petals, and conversion of sepals into carpel-like structures that grew into fruits and ripened. Our findings suggest that LeHB-1 is not only involved in the control of ripening but also plays a critical role in floral organogenesis.

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          A naturally occurring epigenetic mutation in a gene encoding an SBP-box transcription factor inhibits tomato fruit ripening.

          A major component in the regulatory network controlling fruit ripening is likely to be the gene at the tomato Colorless non-ripening (Cnr) locus. The Cnr mutation results in colorless fruits with a substantial loss of cell-to-cell adhesion. The nature of the mutation and the identity of the Cnr gene were previously unknown. Using positional cloning and virus-induced gene silencing, here we demonstrate that an SBP-box (SQUAMOSA promoter binding protein-like) gene resides at the Cnr locus. Furthermore, the Cnr phenotype results from a spontaneous epigenetic change in the SBP-box promoter. The discovery that Cnr is an epimutation was unexpected, as very few spontaneous epimutations have been described in plants. This study demonstrates that an SBP-box gene is critical for normal ripening and highlights the likely importance of epialleles in plant development and the generation of natural variation.
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            The major clades of MADS-box genes and their role in the development and evolution of flowering plants.

            A. Becker (2003)
            MADS-box genes encode a family of transcription factors which control diverse developmental processes in flowering plants ranging from root to flower and fruit development. Sequencing of (almost) the complete Arabidopsis genome enabled the identification of (almost) all of the Arabidopsis MADS-box genes. MADS-box genes have been divided in two large groups, termed type I and type II genes. The type II genes comprise the MEF2-like genes of animals and fungi and the MIKC-type genes of plants. The majority of MIKC-type genes are of the MIKC(c)-type, which includes all plant MADS-box genes for which expression patterns or mutant phenotypes are known. By phylogeny reconstruction, almost all of the MIKC(c)-type genes can be subdivided into 12 major gene clades, each clade comprising 1-6 paralogs from Arabidopsis and putative orthologs from other seed plants. Here we first briefly describe the deep branching of the MADS-box gene tree to place the MIKC(c)-type genes into an evolutionary context. For every clade of MIKC(c)-type genes we then review what is known about its members from Arabidopsis and well-studied members from other phylogenetically informative plant species. By gene sampling and phylogeny reconstructions we provide minimal estimates for the ages of the different clades. It turns out that 7 of the 12 major gene clades, i.e., AG-, AGL6-, AGL12-, DEF+GLO- (B), GGM13- (B(s)), STMADS11- and TM3-like genes very likely existed already in the most recent common ancestor of angiosperms and gymnosperms about 300MYA. Three of the other clades, i.e., AGL2-, AGL17-, and SQUA-like genes, existed at least already in the most recent common ancestor of monocots and eudicots about 200 MYA. Only for two gene clades, AGL15-like genes (2 genes in Arabidopsis) and FLC-like genes (6 genes) members from plants other than Brassicaceae have not been reported yet. Similarly, only one ancient clade known from other flowering plant species, TM8-like genes, is not represented in Arabidopsis. These findings reveal that the diversity of MADS-box genes in Arabidopsis is rather ancient and representative for other flowering plants. Our studies may thus help to predict the set of MADS-box genes in all other flowering plants, except for relatively young paralogs. For the different gene clades we try to identify ancestral and derived gene functions and review the importance of these clades for seed plant development and evolution. We put special emphasis on gene clades for which insights into their importance has rapidly increased just recently.
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              MOLECULAR BIOLOGY OF FRUIT MATURATION AND RIPENING.

              The development and maturation of fruits has received considerable scientific scrutiny because of both the uniqueness of such processes to the biology of plants and the importance of fruit as a significant component of the human diet. Molecular and genetic analysis of fruit development, and especially ripening of fleshy fruits, has resulted in significant gains in knowledge over recent years. Great strides have been made in the areas of ethylene biosynthesis and response, cell wall metabolism, and environmental factors, such as light, that impact ripening. Discoveries made in Arabidopsis in terms of general mechanisms for signal transduction, in addition to specific mechanisms of carpel development, have assisted discovery in more traditional models such as tomato. This review attempts to coalesce recent findings in the areas of fruit development and ripening.
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                Author and article information

                Journal
                Plant J
                tpj
                The Plant Journal
                Blackwell Publishing Ltd
                0960-7412
                1365-313X
                July 2008
                27 May 2008
                : 55
                : 2
                : 301-310
                Affiliations
                [1 ]Plant Sciences Division, School of Biosciences, University of Nottingham Sutton Bonington Campus, Loughborough, LE12 5RD, UK
                [2 ]Warwick HRI, University of Warwick Wellesbourne, Warwick CV35 9EF, UK,
                [3 ]College of Horticulture, Northwest A & F University Yangling, Shaanxi 712100, China
                Author notes
                *For correspondence (fax +44 0 1159516334; e-mail don.grierson@ 123456nottingham.ac.uk )
                Article
                10.1111/j.1365-313X.2008.03505.x
                2607530
                18397374
                f2441542-13a0-4432-9de5-9de6ca8a81b0
                © 2008 The Authors Journal compilation © 2008 Blackwell Publishing Ltd

                Re-use of this article is permitted in accordance with the Creative Commons Deed, Attribution 2.5, which does not permit commercial exploitation.

                History
                : 24 January 2008
                : 11 March 2008
                : 17 March 2008
                Categories
                Original Articles

                Plant science & Botany
                tomato,leaco1,floral organogenesis,lehb-1,hd-zip homeobox protein,ripening
                Plant science & Botany
                tomato, leaco1, floral organogenesis, lehb-1, hd-zip homeobox protein, ripening

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