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      The MADS and the Beauty: Genes Involved in the Development of Orchid Flowers

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          Abstract

          Since the time of Darwin, biologists have studied the origin and evolution of the Orchidaceae, one of the largest families of flowering plants. In the last two decades, the extreme diversity and specialization of floral morphology and the uncoupled rate of morphological and molecular evolution that have been observed in some orchid species have spurred interest in the study of the genes involved in flower development in this plant family. As part of the complex network of regulatory genes driving the formation of flower organs, the MADS-box represents the most studied gene family, both from functional and evolutionary perspectives. Despite the absence of a published genome for orchids, comparative genetic analyses are clarifying the functional role and the evolutionary pattern of the MADS-box genes in orchids. Various evolutionary forces act on the MADS-box genes in orchids, such as diffuse purifying selection and the relaxation of selective constraints, which sometimes reveals a heterogeneous selective pattern of the coding and non-coding regions. The emerging theory regarding the evolution of floral diversity in orchids proposes that the diversification of the orchid perianth was a consequence of duplication events and changes in the regulatory regions of the MADS-box genes, followed by sub- and neo-functionalization. This specific developmental-genetic code is termed the “orchid code.”

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          Complexes of MADS-box proteins are sufficient to convert leaves into floral organs.

          T Honma, K Goto (2001)
          Genetic studies, using floral homeotic mutants, have led to the ABC model of flower development. This model proposes that the combinatorial action of three sets of genes, the A, B and C function genes, specify the four floral organs (sepals, petals, stamens and carpels) in the concentric floral whorls. However, attempts to convert vegetative organs into floral organs by altering the expression of ABC genes have been unsuccessful. Here we show that the class B proteins of Arabidopsis, PISTILLATA (PI) and APETALA3 (AP3), interact with APETALA1 (AP1, a class A protein) and SEPALLATA3 (SEP3, previously AGL9), and with AGAMOUS (AG, a class C protein) through SEP3. We also show that vegetative leaves of triply transgenic plants, 35S::PI;35S::AP3;35S::AP1 or 35S::PI;35S::AP3;35S::SEP3, are transformed into petaloid organs and that those of 35S::PI;35S::AP3;35S::SEP3;35S::AG are transformed into staminoid organs. Our findings indicate that the formation of ternary and quaternary complexes of ABC proteins may be the molecular basis of the ABC model, and that the flower-specific expression of SEP3 restricts the action of the ABC genes to the flower.
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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 characterization of the Arabidopsis floral homeotic gene APETALA1.

              The first step in flower development is the transition of an inflorescence meristem into a floral meristem. Each floral meristem differentiates into a flower consisting of four organ types that occupy precisely defined positions within four concentric whorls. Genetic studies in Arabidopsis thaliana and Antirrhinum majus have identified early-acting genes that determine the identify of the floral meristem, and late-acting genes that determine floral organ identity. In Arabidopsis, at least two genes, APETALA1 and LEAFY, are required for the transition of an influorescence meristem into a floral meristem. We have cloned the APETALA1 gene and here we show that it encodes a putative transcription factor that contains a MADS-domain. APETALA1 RNA is uniformly expressed in young flower primordia, and later becomes localized to sepals and petals. Our results suggest that APETALA1 acts locally to specify the identity of the floral meristem, and to determine sepal and petal development.
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                Author and article information

                Journal
                Curr Genomics
                CG
                Current Genomics
                Bentham Science Publishers Ltd
                1389-2029
                1875-5488
                August 2011
                : 12
                : 5
                : 342-356
                Affiliations
                []Department of Biological Sciences, University of Naples Federico II, Via Mezzocannone 8, 80134 Napoli, Italy
                Author notes
                [* ]Address correspondence to this author at the Department of Biological Sciences, University of Naples Federico II, Via Mezzocannone 8, 80134 Napoli, Italy; Tel: +390812535190; Fax: +390812535015; E-mail: serena.aceto@ 123456unina.it
                Article
                CG-12-342
                10.2174/138920211796429754
                3145264
                22294877
                5c2c9d30-2171-4b08-90ee-2d0c0824764b
                ©2011 Bentham Science Publishers Ltd.

                This is an open access article distributed under the terms of the Creative Commons Attribution License ( http://creativecommons.org/licenses/by/2.5/), which permits unrestrictive use, distribution, and reproduction in any medium, provided the original work is properly cited.

                History
                : 5 April 2011
                : 16 June 2011
                : 21 June 2011
                Categories
                Article

                Genetics
                orchid code.,flower development,orchidaceae,mads-box genes
                Genetics
                orchid code., flower development, orchidaceae, mads-box genes

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