The pollen-collecting hairs of Campanula (Campanulaceae). I. Morphological variation and the retractive mechanism

The pollen-collecting hairs (PCHs) of Campanula have been a subject of intense debate for the past two centuries. Although several morphological studies have been made on these hairs, detailed comparative studies among species are still lacking, their function and adaptive significance being an unsolved question. The present study comprises two microscopy techniques: scanning electron microscopy and confocal scanning laser microscopy. The aim of the present study is to elucidate: 1) the variation in morphology of the PCHs, 2) the variation in presence/absence of the PCHs by the time of spreading of the stigmatic lobes, 3) the variation in the retractive mechanism of the PCHs, and 4) the correlation between pollination and the retraction of the PCHs. In several species PCHs of various lengths are found. Despite the variations in length of the hairs, the same retractive mechanism is found in all species studied. In most species the hairs retract into basal cavities within the style late in anthesis. The cells into which the hairs retract differ in length among species. Pollen grains are often found within the cavities together with the retracted hairs, a mechanism considered to prevent self-pollination. Pollen germination within the cavities was not observed. In a few species, the PCHs are still present at stigma receptivity. Differences in the shape and size of the cells surrounding the PCHs are documented. The diameter of the pits and the pollen grains vary among species. Other types of hairs on the style are recognized in some species, being of various lengths. These other types do not retract at stigma development and should not be regarded as pollen-collectors. They possibly facilitate for visiting insects to reach the nectar glands, present at the top of the ovary.