At maturity, the pollen of Lilium spp. is coated with an adhesive, pigmented material, the Pollenkitt of German authors. This material contains a lipid component and carotenoid pigments; in L. longiflorum the principal pigment is α-carotene 5,6-epoxide. Synthesis of the coat materials takes place in the tapetum. The pigments accumulate in osmiophilic globuli from the time of release of the spores from the tetrad until the dissolution of the tapetum as a tissue. The lipids and associated pigments are then discharged into the loculi, where they migrate between the spores, filling the recesses in the surface of the exine.