Male/Female Trade-Off in Hermaphroditic Y-Chromosome Deletion Mutants of the Dioecious Plant Silene latifolia

The evolution of dioecy from hermaphroditism allows for avoidance of self-pollination, and its evolutionary background has been investigated both experimentally and theoretically since it was first proposed by Darwin. To reproduce this evolution, we screened hermaphroditic mutants of Silene latifolia using heavy-ion beam or gamma-ray irradiation and characterized the phenotypes of their floral organs. Our scatterplots indicate severe deviations from the trade-off relationships between pollen and ovule numbers and between seed and germinated pollen numbers in hermaphroditic mutant S. latifolia. These deviations presumably led to promotion of dioecy from the ancestral state of S. latifolia. To infer the likely flower phenotypic characteristics of the ancestral plant of S. latifolia before evolving dioecy, the flowers of Silene viscosa, a naturally hermaphroditic plant related to S. latifolia were also characterized. S. viscosa exhibits both spatial separation of stamens from pistils within the flower (reverse herkogamy) and temporal separation of stamen and pistil maturation (dichogamy), raising the question of whether hermaphroditic mutant S. latifolia, which is thought to be the ancestral state, would possess these functions. We show that two hermaphroditic mutants of the dioecious plant S. latifolia exhibit signs of protogyny (reverse dichogamy) and approach herkogamy, as pistils were constantly longer than stamens. These findings illustrate the evolution of dioecy from hermaphroditism as a self-pollination avoidance mechanism and to balance the investments into male and female functions.