from this damage as well as a physically robust container. Analysis of sporopollenin from the clubmoss Lycopodium in the late 1980s have shown distinct structural differences from that of flowering plants. In 2020, more detailed analysis of sporopollenin from Lycopodium clavatum provided more structural information. It showed a complete lack of aromatic structures and the presence of a macrocyclic backbone of polyhydroxylated tetraketide-like monomers with pseudo-aromatic 2-pyrone rings. These were crosslinked to a poly(hydroxy acid) chain by ether linkages to form the polymer. Biosynthesis Electron microscopy shows that the tapetal cells that surround the developing pollen grain in the anther have a highly active secretory system containing lipophilic globules. These globules are believed to contain sporopollenin precursors. Tracer experiments have shown that phenylalanine is a major precursor, but other carbon sources also contribute. The biosynthetic pathway for phenylpropanoid is very active in tapetal cells, supporting the idea that its products are needed for sporopollenin synthesis. Chemical inhibitors of pollen development and many male sterile mutants have effects on the secretion of these globules by the tapetal cells. See also Chitin Conchiolin Tectin References Biomolecules Organic polymers Pollination Palynology