The morphological relationship between carpels and ovules in angiosperms: pitfalls of morphological interpretation

Abstract

Carpels and ovules have been differently interpreted over the past two centuries. In this review, some of these interpretations are highlighted, with particular emphasis on the current situation. Ovules are part of and are enclosed in carpels in all living angiosperms. Living angiosperms are monophyletic, and the evolutionary association between ovules and the leaf-like part, the carpel wall, had taken place at or before the time the clade of extant angiosperms was established. From what we know at present, there are no cauline' ovules in extant angiosperms. Developmentally, carpel walls and ovules are not always synchronous across all extant angiosperms. In early development ovules may be relatively precocious or relatively late compared with carpel walls. They are late in early-diverging angiosperms (ANITA grade, magnoliids, some early-diverging eudicots) but precocious in some more derived groups (e.g. some Caryophyllales and Primulaceae). Carpel primordia have a certain depth in the floral apex, and the entire activated area of a carpel primordium may be several cell layers thick. Thus, the carpel is embedded' or rooted' within the remaining floral apex. The parts of a carpel develop at different times in carpel ontogeny and probably evolved at different times on the line leading to the angiosperms, which needs to be considered in interpretations. Carpel development depends on a complex genetic network, which increased stepwise over evolutionary time and contains hundreds of genes revealed in molecular developmental biology. The evolutionary history of such networks in carpel walls and ovules is unlikely to be easily disentangled, as most of these genes are not transcription factors.