The biosynthesis, composition and assembly of the outer pollen wall: A tough case to crack.

The formation of the durable outer pollen wall, largely composed of sporopollenin, is essential for the protection of the male gametophyte and plant reproduction. Despite its apparent strict conservation amongst land plants, the composition of sporopollenin and the biosynthetic pathway(s) yielding this recalcitrant biopolymer remain elusive. Recent molecular genetic studies in Arabidopsis thaliana (Arabidopsis) and rice have, however, identified key genes involved in sporopollenin formation, allowing a better understanding of the biochemistry and cell biology underlying sporopollenin biosynthesis and pollen wall development. Herein, current knowledge of the biochemical composition of the outer pollen wall is reviewed, with an emphasis on enzymes with characterized biochemical activities in sporopollenin and pollen coat biosynthesis. The tapetum, which forms the innermost sporophytic cell layer of the anther and envelops developing pollen, plays an essential role in sporopollenin and pollen coat formation. Recent studies show that several tapetum-expressed genes encode enzymes that metabolize fatty acid derived compounds to form putative sporopollenin precursors, including tetraketides derived from fatty acyl-CoA starter molecules, but analysis of mutants defective in pollen wall development indicate that other components are also incorporated into sporopollenin. Also highlighted are the many uncertainties remaining in the development of a sporopollenin-fortified pollen wall, particularly in relation to the mechanisms of sporopollenin precursor transport and assembly into the patterned form of the pollen wall. A working model for sporopollenin biosynthesis is proposed based on the data obtained largely from studies of Arabidopsis, and future challenges to complete our understanding of pollen wall biology are outlined.