Genes Controlling Flower Development in Plants

The Steps of Flower Development - Genes Are Implicated

Mendelian Genes Define the Committment to Flowering

Mendelian Genes Define Floral Organ Identity

Cloning Committment to Flowering and Flower Organ Genes

MADS-Box Genes

Analyzing Gene Expression with In Situ Hybridization

The Molecular Expression of Floral Committment Genes

The Molecular Expression of Floral Organ Genes

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Mendelian Genes Define Floral Organ Identity

A model was developed that attempts to explain the interactions of the different genes that control floral organ identity. All the genes are classified as expressing one of three activities, A, B, or C. The A gene activities control the development of the sepal and petal, B gene activities control petal and stamen development, and C gene activities control stamen and carpel development. The following diagram depicts these interactions.

The two A function genes are APETALA2 and APETALA1. Alleles of these two genes have been isolated that show varying degrees of effect, but in general if an A function gene is mutated, the first whorl develops as a carpelc and the second whorl develops as a stamen. ovulata is an A function gene of snapdragon similar to APETALA2. The following image shows the APETALA2 mutant.

The B gene functions are defined by the genes APETALA3 and PISTILLATA. The net effect of B gene mutations is that whorl 2 develops as a sepal rather than a petal, and whorl 3 develops as a carpel not a stamen. deficiens and globosa are snapdragon genes that have homologous functions to the Arabidopsis B function genes. The following image shows the APETALA3 mutant.

Finally, C gene functions are defined by the gene AGAMOUS. Mutants of this gene have the third whorl stamen replaced by a petal, and the fourth whorl develops into a new flower with the sepal-petal-petal pattern. Furthermore, flower development in AGAMOUS mutants is indeterminate, not determinate. A snapdragon gene similar to AGAMOUS is pleniflora. The following image shows the AGAMOUS mutant.

The following table summaries the phenotypic effects of mutations in each of the three functions.

**Phenotypic Effects of Mutations in A, B or C Function Floral Identity Genes**
	Phenotype
Mutation	Whorl 1	Whorl 2	Whorl 3	Whorl 4
Wild Type	Sepal	Petal	Stamen	Carpel
A Function	Carpel	Stamen	Stamen	Carpel
B Function	Sepal	Sepal	Carpel	Carpel
C Function	Sepal	Petal	Petal	New Flower

Several conclusions can be drawn regarding the functions of these genes by studying single and double mutants. Because a mutation of an A function gene results in the expressiion of organ phenotypes controlled by C function genes, it appears that A gene functions repress the expression of the C gene functions in the whorls giving rise to sepals and petals. Likewise, the appearance of the petal in the third whorl of C gene mutants, suggest that C genes repress the activities of A genes in the organs that they control.

These conclusions are based on single mutants. For example, APETALA2 A function mutants develop C function organs, carpels and stamens, in the first two whorls, respectively. What would be expected from an A and C double mutant? This mutant would not be expected to have any functions exclusively controlled by the the A and C function genes. And indeed this is what was seen when the APETALA2/AGAMOUS double mutant was developed. The first whorl develops as a leaf and the second whorl has stamen-like petals. This second whorl phenotype of this mutant is the result of the activities of the B gene functions.

Finally what would be expected with an A, B and C function triple mutant? This mutant would have no genes functioning that determine normal floral organ development. As expected, the triple mutants lack any floral organs, and the flower essentially consists of leaves developing form each of the whorls.