A keystone-like grain in hornet comb cells: Its nature and physical properties

In the roof of cells built by the Oriental hornet, Vespa orientalis (Vespinae, Hymenoptera), there is one or more grains differing in their composition and shape from the surrounding structures. These grains have a diameter of 100-200 mum and protrude slightly from the roof inward. They contain primarily a large percentage of Ti but also Fe and occasionally Si. The mineral makeup in the grains is still uncertain, but it seems that the minerals belong to the group called perovskite. These grains in the hornet's comb may originate from the soil around the nest, that is, comprised of ceramic matter, but it is also possible that they are secreted from a gland within the frons plate of the hornet because the fungus-like secretion in this gland contains elements similar to those in the roof of the comb cells. We conjecture that the grain in question serves as a reference point for the hornets that build the cell walls in the direction of gravity. The cement whereby the building hornets conjoin the various building materials, whether organic (cellulose and the like) or inorganic (sand grains or tiny stones), is a saliva that is secreted from their oral gland, which upon release hardens into a polymeric glue. The ceramic grain embedded within this polymeric matrix is endowed with different physical properties and can therefore respond differently to the tension or pressure of the surrounding walls.