Annulus-pore relationship in Gramineae (Poaceae) pollen: the pore margin of Pariana.

Pariana, a primitive bamboo, is the only genus in the Gramineae (Poaceae) to have pollen grains without an annulus as part of its single aperture (porate) system. In contrast, the markedly thickened exine layer underlying the pore margin is similar to counterparts in all grass genera. Components of the future annulus in Gramineae pollen develop toward the cytoplasm (proximally) and begin to be pressed outward by an increase in the cytoplasm during the microspore vacuolate stage, culminating in an annulus by maturity. However, in some species of Pariana these components are either not sufficiently developed or the cytoplasmic expansion is not sufficient to press the components into an annular ring around the pore. The structural relationship of exine layering in this type of pollen grain in Gramineae and other families with similar apertures has not hitherto been extensively studied. A critical examination of the apertures in bambusoid grasses may clarify their systematic position within the Gramineae.