Apomixis in flowering plants: an overview.

Apomixis is a common feature of perennial plants, which occurs in ca. 60% of the British flora, but has been largely ignored by reproductive theoreticians. Successful individuals may cover huge areas, and live to great ages, favoured by 'symmetrical' selection. Apomixis is favoured by colonizing modes, for instance post-glacially. Despite its theoretical advantages, apomixis usually coexists with sexuality, suggesting 'hidden' disadvantages. Agamospermy (apomixis by seed) is relatively uncommon, but gains from the attributes of the seed. It pays agamospermy genes, which discourage recombination, to form co-adapted linkage groups, so that they become targets for disadvantageous recessive mutant accumulation. Consequently, agamospermy genes cannot succeed in diploids and agamosperms are hybrid and highly heterotic. Agamospermous endosperm may suffer from genomic imbalance, so that nutritious ovules, which can support embryos without endosperm, may be preadapted for agamospermy. When primary endosperm nucleus fertilization ('pseudogamy') continues as a requirement for many aposporous agamosperms, selfing sex becomes preadaptive and archesporial sex remains an option. Apomictic populations can be quite variable although apomictic families are much less variable than sexuals. Only in some diplosporous species does sex disappear completely, and in those species some release of variability may persist through somatic recombination. The search for an agamospermy gene suitable for genetic modification should target fertile sexuals with a single localized agamospermy (A) gene, which therefore lack a genetic load. The A gene should coexist alongside sexuality, so that it would be easy to select seedlings of sexual and asexual origins. Plants with sporophytic agamospermy provide all these attributes.