Hepburn summarizes what was known in 2014

> What we already know from these various experiments and observations is that a dietary intake of pollen, equated with protein, is essential for the normal development of the wax glands. Once developed, the glands may well function without additional pollen.   

> The findings of Taranov (1959) that the influx of fresh, field pollen into a colony stimulates wax production, hence comb construction, and are extremely difficult to interpret because the variables are conflated and inevitably coupled to fine weather, suitable ambient temperatures, and presumably the greater activities of the foragers. The task of unraveling the properties of these obviously complex stimuli remains as yet to be done.

> The discovery that beeswax contains proteinaceous material (Kurstjens et a1. 1985) also supports a fundamental role of protein in beeswax synthesis, that extending well beyond what is essential for the development of the cytoplasmic vacuolar system of the wax gland cells.

Pete B

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