Late-acting self-incompatibility--the pariah breeding system in flowering plants

It is estimated that around half of all species of flowering plants show self-incompatibility (SI). However, the great majority of species alleged to have SI simply comply with 'the inability of a fully fertile hermaphrodite plant to produce zygotes when self-pollinated'--a definition that is neutral as to cause. Surprisingly few species have been investigated experimentally to determine whether their SI has the type of genetic control found in one of the three established mechanisms, that is, homomorphic gametophytic, homomorphic sporophytic or heteromorphic SI. Furthermore, our knowledge of the molecular basis of homomorphic SI derives from a few species in just five families--a small sample that has nevertheless revealed the existence of three different molecular mechanisms. Importantly, a sizeable cohort of species are self-sterile despite the fact that self-pollen tubes reach the ovary and in most cases penetrate ovules, a phenomenon called late-acting self-incompatibility (LSI). This review draws attention to the confusion between species that show 'self-incompatibility' and those that possess one of the 'conventional SI mechanisms' and to argue the case for recognition of LSI as having a widespread occurrence and as a mechanism that inhibits selfing and promotes outbreeding in many plant species.