Gynoecium structure and development in core Caryophyllales: a matter of proportions

Abstract

Abstract This review based on a morphological and developmental perspective reveals a striking diversity in shapes and evolutionary trends in the gynoecium of core Caryophyllales that have affected the number of carpels, the formation of septa and the number of ovules. Two major developmental shifts are responsible for the diversity in gynoecial forms and are linked to the proportional development of carpellary tissue (ovary wall) and the floral apex. (1) Meristic change is caused by an expansion or reduction of the diameter of the floral apex. An expansion leads to polygyny linked with the development of more numerous small carpels; a reduction of space leads to lower carpel numbers, eventually resulting in a single carpel. (2) Different ovary shapes can be recognized at a mid-developmental stage predicting the further development of ovaries. With an equal growth of the ovary wall and floral apex, young ovaries take the shape of a salt-shaker; with more extensive development of the floral apex and delay of the ovary wall, a club-shaped ovary is formed; with faster growth of the ovary wall linked with intercalary expansion and a delayed growth of the floral apex, a (half-) inferior cup-shaped ovary develops. The different growth forms are the results of heterochronic shifts and affect the development of septa and ovule numbers. A common trend in the order implies a weakening and break-up of septa during development, leading to residual apical and basal septa and the shift to free-central and basal placentation. The club-shaped ovary is linked with an almost complete loss of septa and a reduction of the ovule number to one. The salt-shaker shape leads to ovaries with a massive placental column and several ovules. The cup-shaped ovary leads to a shift of ovules away from the floral apex. Developmental flexibility is responsible for a disconnection of carpel wall growth from ovular tissue. Subtle shifts in proportional growth lead to a high diversification of ovaries in core Caryophyllales and the establishment of predictable developmental trends. These trends clearly represent apomorphic tendencies, affecting different families of core Caryophyllales in different degrees. The ancestral gynoecium was probably pentamerous and isomerous with the other floral whorls, with ovules clearly separated from the carpellary wall and inserted on axile placentas corresponding to the central axis of the flower.