Reproductive versatility and the epigenetic control of female gametogenesis

Each year, plants and animals perform the task of repopulating the planet through patterns of courtship and mating that have a unifying and compelling logic: the production of offspring. Although life of nearly all organisms is organized around sex and breeding, Darwinian thinking focused more on the struggle for existence than on evolutionary significance of this frantic race to reproduce. In Darwin's own words, "We do not know the final cause of sexuality; why new beings should be produced by the union of the two sexual elements. The whole subject is hidden in darkness…" (Darwin 1862). In plants, a major consequence of this search for survival is the evolution of a multitude of reproductive alternatives that have intrigued botanists, geneticists, and evolutionary biologists for more than 100 years. Because sexually derived genetic diversity is interpreted as essential for adaptation, it is often thought that sex is necessary for the perpetuation of a species; however, many organisms--including several hundred families of flowering plants--are going efficiently about propagating their kind without bothering with meiosis and mating. Whereas many plants can undergo vegetative propagation, through the production of stolons, bulbs, or rhizomes, for example, many others have developed methods to produce an embryo from a single cell whose nucleus is not formed by the fusion of two gametes, offering a direct developmental and evolutionary challenge to sexual reproduction. Recent evidence suggests that epigenetic mechanisms that control transcriptional silencing of DNA repetitive elements and heterochromatin are crucial for the acquisition of cell identity in the ovule, opening the possibility that the developmental distinction between sexual development and apomixis might have evolved as an adaptive response to evolutionary forces that modulate structural variation and reproductive versatility in flowering plants.