JXB Advance Access originally published online on April 18, 2005
Journal of Experimental Botany 2005 56(416):1665-1674; doi:10.1093/jxb/eri163
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RESEARCH PAPER |
Phosphoinositides and phosphatidic acid regulate pollen tube growth and reorientation through modulation of [Ca2+]c and membrane secretion
1Universidade de Lisboa, Faculdade de Ciências de Lisboa, ICAT, 1749-016 Lisboa, Portugal
2Institute of General Botany, University of Hamburg, Ohnhorst-Straße 18, D-22609 Hamburg, Germany
3University of Hannover, Institut für Zierpflanzenanbau, Baumschule u. Pflanzenzüchtung, Herrenhäuser Str. 2, D-30419 Hannover, Germany
To whom correspondence should be addressed. Fax: +351 217 500 048. E-mail: r.malho{at}fc.ul.pt
The maintenance of a calcium gradient and vesicle secretion in the apex of pollen tubes is essential for growth. It is shown here that phosphatidylinositol-4,5-bisphosphate (PIP2) and D-myo-inositol-1,4,5-trisphosphate (IP3), together with phosphatidic acid (PA), play a vital role in the regulation of these processes. Changes in the intracellular concentration of both PIP2 and IP3 (induced by photolysis of caged-probes), modified growth and caused reorientation of the growth axis. However, measurements of cytosolic free calcium ([Ca2+]c) and apical secretion revealed significant differences between the photorelease of PIP2 or IP3. When released in the first 50 µm of the pollen tube, PIP2 led to transient growth perturbation, [Ca2+]c increases, and inhibition of apical secretion. By contrast, a concentration of IP3 which caused a [Ca2+]c transient of similar magnitude, stimulated apical secretion and caused severe growth perturbation. Furthermore, the [Ca2+]c transient induced by IP3 was spatially different causing a pronounced elevation in the sub-apical region. These observations suggest different targets for the two phosphoinositides. One of the targets is suggested to be PA, a product of PIP2 hydrolysis via phospholipase C (PLC) or phospholipase D (PLD) activity. It was found that antagonists of PA accumulation (e.g. butan-1-ol) and inhibitors of PLC and PLD reversibly halted polarity. Reduction of PA levels caused the dissipation of the [Ca2+]c gradient and inhibited apical plasma membrane recycling. It was also found to cause abolition of the apical zonation. These data suggest that phosphoinositides and phospholipids regulate tip growth through a multiple pathway system involving regulation of [Ca2+]c levels, endo/exocytosis, and vesicular trafficking.
Key words: Ins(1,4,5)P3, phosphatidic acid, phospholipases, PIP2, secretion
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