The discovery of the MADS-box genes and the study of model plants such as Arabidopsis thaliana and Antirrhinum majus have greatly improved our understanding of the molecular mechanisms driving the diversity in floral development. The class B genes, which belong to the MADS-box gene family, are important regulators of the development of petals and stamens in flowering plants. Many nongrass monocot flowers have two whorls of petaloid organs, which are called tepals. To explain this floral morphology, the modified ABC model was proposed. This model was exemplified by the tulip, in which expansion and restriction of class B gene expression is linked to the transition of floral morphologies in whorl 1. The expression patterns of class B genes from many monocot species nicely fit this model; however, those from some species, such as asparagus, do not. In this review, we summarize the relationship between class B gene expression and floral morphology in nongrass monocots, such as Liliales (Liliaceae) and Asparagales species, and discuss the applicability of the modified ABC model to monocot flowers.

B class MADS-box genes play important roles in petal and stamen development. Some monocotyledonous species, including liliaceous ones, produce flowers with petaloid tepals in whorls 1 and 2. A modified ABCE model has been proposed to explain the molecular mechanism of development of two-layered petaloid tepals. However, direct evidence for this modified ABCE model has not been reported to date. To clarify the molecular mechanism determining the organ identity of two-layered petaloid tepals, we used chimeric repressor gene-silencing technology (CRES-T) to examine the suppression of B function in the liliaceous ornamental Tricyrtis sp. Transgenic plants with suppressed B class genes produced sepaloid tepals in whorls 1 and 2 instead of the petaloid tepals as expected. In addition, the stamens of transgenic plants converted into pistil-like organs with ovule- and stigma-like structures. This report is the first to describe the successful suppression of B function in monocotyledonous species with two-layered petaloid tepals, and the results strongly support the modified ABCE model.

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Application of forchlorfenuron (CPPU) to flower buds induces morphologically different paracorollas, i.e., wide and narrow type, in torenia (Torenia fournieri L.). We investigated the morphological properties and the role of floral homeotic genes in the formation of these two types of paracorolla. The morphology of epidermal cells and distribution pattern of vascular bundles in the wide paracorolla was the same as in the petal; however, in the narrow paracorolla, the morphology of epidermal cells was either petal-like or stamen-like, and the distribution pattern of vascular bundles was stamen-like. In situ hybridization analysis of the floral homeotic genes showed that a class A gene, T. fournieri SQUAMOSA (TfSQUA), and class B genes, TfDEFICIENS (TfDEF) and TfGLOBOSA (TfGLO), were expressed in the broad part of the primordia of the wide paracorolla, as in the petal. Class C genes, TfPLENA1 (TfPLE1) and TfFARINELLI (TfFAR), were only expressed at the margin of the primordia. However, in the primordia of the narrow paracorolla, TfSQUA was only expressed at the margin, but the class B genes and one of the class C genes (TfPLE1) were expressed in a broad section, as in the primordia of the wide paracorolla. This expression pattern in the narrow paracorolla was intermediate between that of the petal and the stamen. In later developmental stages, quantitative real-time PCR analyses showed that, in the wide paracorolla, TfSQUA and class B genes were highly expressed but the expression of class C genes, as in the petal, was low. In the narrow paracorolla, class B genes were also highly expressed as in the petal; however, the expression of both TfSQUA and the class C genes was low, as in the stamen and the petal, respectively. This expression pattern probably reflects the unstable floral organ identity of the narrow paracorolla, and the expression pattern in paracorollas is determined by the site where the paracorolla is formed.

According to the modified ABC model, class B genes are expressed in whorl 1 as well as in whorls 2 and 3, therefore the organs of whorls 1 and 2 have the same petaloid structure in many monocots. The viridiflora tulip, a well-known cultivar group, has flowers in which the tepals in whorls 1 and 2 have greenish stripes. This phenotype could be expected as class B mutant by the modified ABC model. In this study, we discovered that tepal phenotypes correlated with that of stamens. We isolated two class A genes (TGSQA and TGSQB) from wild type tulip. Northern hybridization of these two class A genes, and the class B genes (TGDEFA, TGDEFB and TGGLO), showed that the expression of TGDEFA and TGDEFB in the viridiflora tulips were weaker than those in wild type cultivars, whereas the expression patterns of the rest genes were almost identical. From our results, we suggest that reduced expression of the two DEF-like genes, TGDEFA and TGDEFB, is involved in the development of the viridiflora phenotype. This reduced expression could be caused by the amino acid difference.

Alstroemeria (Liliales) has two layers of petaloid tepals, in which the often spotted narrow inner tepals can be distinguished easily from the wider outer tepals. In order to explore this floral morphology in Alstroemeria, we investigated the tepal morphology and the expression patterns of three class B genes, whose homologs in eudicots have been shown previously to be involved in petal and stamen development. The two DEF-like genes (AlsDEFa and AlsDEFb) and the one GLO-like gene (AlsGLO) of Alstroemeria ligtu were isolated by rapid amplification of cDNA ends (RACE). Northern hybridization, reverse transcription-PCR (RT-PCR) and in situ hybridization analyses indicated that AlsDEFb and AlsGLO were expressed in whorls 1, 2 and 3 (outer tepals, inner tepals and stamens, respectively), whereas AlsDEFa expression was detected only in whorls 2 and 3. These results suggest that in A. ligtu, AlsDEFb and AlsGLO would participate in determining the organ identity of the two-layered petaloid tepals and stamens, which is in support of the modified ABC model. Additionally, the distinctive expression patterns of AlsDEFa and AlsDEFb might be related to morphological differences between the two-layered tepals.

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