Modulation of endocytosis in pollen tube growth by phosphoinositides and phospholipids

In plants, tip-growing cells represent an ideal system to investigate signal transduction mechanisms, and among those, pollen tubes are one of the favourite models. Many signalling pathways have been identified during germination and tip growth, namely, Ca2+, calmodulin, phosphoinositides, cyclic AMP, and GTPases. Not surprisingly, the apical secretory machinery, essential for tip growth, seems to be an intersection point for all these pathways. Recently, the phospholipid phosphatidic acid was also suggested to actively participate in the control of endo- and exocytosis and to interfere with the correct positioning of the actin cytoskeleton. Phosphatidic acid seems to act concertedly with the phosphoinositides phosphatidylinositol 4,5-bisphosphate and D-myo-inositol 1,4,5-trisphosphate. Here we review previous data and discuss additional evidence that these three molecules have a combined action modulating both the actin cytoskeleton and the apical secretory machinery. We further discuss how these findings can be integrated into a working model for pollen tube apical secretion that contemplates the existence of a rapid endocytosis mechanism.