Sandra Doyle

floricaula: A homeotic gene required for flower development in antirrhinum majus

Plants carrying the floricaula (flo) mutation cannot make the transition from inflorescence to floral meristems and have indeterminate shoots in place of flowers. The flo-613 allele carries a Tam3 transposon insertion, which allowed the isolation of the flo locus. The flo gene encodes a putative protein (FLO) containing a proline-rich N-terminus and a highly acidic region. In situ hybridization shows that the flo gene is transiently expressed in the very early stages of flower development. The earliest expression seen is in bract primordia, followed by sepal, petal, and carpel primordia, but no expression is detected in stamen primordia. This pattern of expression has implications for how flo affects phyllotaxis, organ identity, and determinacy. We propose that flo interacts in a sequential manner with other homeotic genes affecting floral organ identity.

Dual role for fimbriata in regulating floral homeotic genes and cell division in Antirrhinum.

The fimbriata (fim) gene of Antirrhinum affects both the identity and arrangement of organs within the flower, and encodes a protein with an F‐box motif. We show that FIM associates with a family of proteins, termed FAPs (FIM‐associated proteins), that are closely related to human and yeast Skp1 proteins. These proteins form complexes with F‐box‐containing partners to promote protein degradation and cell cycle progression. The fap genes are expressed in inflorescence and floral meristems in a pattern that incorporates the domain of fim expression, supporting an in vivo role for a FIM–FAP complex. Analysis of a series of novel fim alleles shows that fim plays a key role in the activation of organ identity genes. In addition, fim acts in the regions between floral organs to specify the correct positioning and maintenance of morphological boundaries. Taking these results together, we propose that FIM–FAP complexes affect both gene expression and cell division, perhaps by promoting selective degradation of regulatory proteins. This may provide a mechanism by which morphological boundaries can be aligned with domains of gene expression during floral development.

Fimbriata controls flower development by mediating between meristem and organ identity genes.

Two major classes of genes directing flower development have so far been described: early activated genes regulating meristem identity and later acting genes controlling organ identity. Here, we show that the fimbriata (fim) gene acts between these two classes in a sequence of gene activation. The fim gene, originally described in 1930, was cloned by transposon tagging from Antirrhinum majus and encodes a product with no detectable homology to other proteins. Mutations in fim result in partial homeotic transformations of floral organs and in reduced determinacy of the meristem. Expression and function of fim depends on the activity of meristem identity genes, and fim in turn controls the spatial and temporal expression of organ identity genes. The pattern of fim expression defines a new domain of the floral meristem that changes with time in a complementary manner to those of the meristem identity gene floricaula and the organ identity gene plena.

Olive: a key gene required for chlorophyll biosynthesis in Antirrhinum majus.

Olive (oli) is a recessive nuclear mutation of Antirrhinum majus which reduces the level of chlorophyll pigmentation and affects the ultrastructure of chloroplasts. The oli‐605 allele carries a Tam3 transposon insertion which has allowed the locus to be isolated. The oli gene encodes a large putative protein of 153 kDa which shows homology to the products of two bacterial genes necessary for tetrapyrrole‐metal chelation during the synthesis of bacteriochlorophyll or cobyrinic acid. We therefore propose that the product of the oli gene is necessary for a key step of chlorophyll synthesis: the chelation of magnesium by protoporphyrin IX. Somatic reversion of the oli‐605 allele produces chimeric plants which indicate that the oli gene functions cell‐autonomously. Expression of oli is restricted to photosynthetic cells and repressed by light, suggesting that it may be involved in regulating the rate of chlorophyll synthesis in green tissues.

Control of flower development and phyllotaxy by meristem identity genes in antirrhinum.

The flower meristem identity genes floricaula (flo) and squamosa (squa) promote a change in phyllotaxy from spiral to whorled in Antirrhinum. To determine how this might be achieved, we have performed a combination of morphological, genetic, and expression analyses. Comparison of the phenotypes and RNA expression patterns of single and double mutants with the wild type showed that flo and squa act together to promote flower development but that flo is epistatic to squa with respect to early effects on phyllotaxy. We propose that a common process underlies the phyllotaxy of wildtype, flo, and squa meristem development but that the relative timing of primordium initiation or growth is altered. This process depends on two separable events: setting aside zones for potential primordium initiation and partitioning these zones into discrete primordia. Failure of the second event can lead to the formation of continuous double spirals, which are occasionally seen in flo mutants.

Homeotic Genes Directing Flower Development in Antirrhinum

Homeotic mutants have been used to define the genetic interactions controlling flowering in Antirrhinum. Three categories of homeotic genes were identified by transposon mutagenesis. The first includes floricaula (flo), which is required to switch inflorescence meristems to floral. This gene has been isolated and shown to be expressed transiently in bract, sepal, petal and carpel primordia. The second group of genes controls the identity (and sometimes the number) of organs in a whorl. These genes affect overlapping whorls and their mutant phenotypes suggest a combinatorial model for gene action in determining the fate of floral primordia. Genes of the third category determine the identity of organs within one whorl and thus affect the symmetry of the flower. We propose that the interactions of these homeotic genes not only control the basic patterns of inflorescence and flower development in Antirrhinum, but possibly in a diverse range of plant species.

Floral Homoeotic and Pigment Mutations Produced by Transposon-Mutagenesis in Antirrhinum majus

Many homoeotic genes affecting flower morphogenesis have been described in diverse species, although there have been few attempts to relate these in a systematic way to the mechanism of floral development (Meyerowitz et al., 1989). In 1742 it was the peloric form of Linaria vulgaris that made Linnaeus change his mind about the fixity of species (Linnaeus, 1744) and over a century later Darwin described, and was intrigued by, the behaviour of the peloric Antirrhinum (Darwin, 1868). We have used transposon-mutagenesis to generate floral homoeotic mutations in Antirrhinum with a view to studying and isolating the genes involved. One advantage of this approach is that transposon integration can be used to identify genes and subsequent excision can be used to prove that the correct gene has been isolated (Martin et al., 1985). In addition, imprecise excision can generate alleles with altered gene expression (Almeida et al., 1989).