Programmed cell death in floral organs: how and why do flowers die?

BACKGROUND: Flowers have a species-specific, limited life span with an irreversible programme of senescence, which is largely independent of environmental factors, unlike leaf senescence, which is much more closely linked with external stimuli. TIMING: Life span of the whole flower is regulated for ecological and energetic reasons, but the death of individual tissues and cells within the flower is co-ordinated at many levels to ensure correct timing. Some floral cells die selectively during organ development, whereas others are retained until the whole organ dies. TRIGGERS: Pollination is an important floral cell death trigger in many species, and its effects are mediated by the plant growth regulator (PGR) ethylene. In some species ethylene is a major regulator of floral senescence, but in others it plays a very minor role and the co-ordinating signals involved remain elusive. Other PGRs such as cytokinin and brassinosteroids are also important but their role is understood only in some specific systems. MECHANISMS: In two floral cell types (the tapetum and the pollen-tube) there is strong evidence for apoptotic-type cell death, similar to that in animal cells. However, in petals there is stronger evidence for an autophagous type of cell death involving endoplasmic reticulum-derived vesicles and the vacuole. Proteases are important, and homologues to animal caspases, key regulators of animal cell death, exist in plants. However, their role is not yet clear. COMPARISON WITH OTHER ORGANS: There are similarities to cell death in other plant organs, and many of the same genes are up-regulated in both leaf and petal senescence; however, there are also important differences for example in the role of PGRs.
CONCLUSIONS: Understanding gene regulation may help to understand cell death in floral organs better, but alone it cannot provide all the answers.