In the past, the way in which bees and other social insects construct comb has been hotly debated. This debate has been ongoing for at least a hundred years



> Thompson (1917) argued that the optimal solution could instead result from physical necessity combined with very rudimentary behaviour. He wrote: ‘the direct efforts of the wasp or bee may be supposed to be limited to the making of a tubular cell, as thin as the nature of the material permits, and packing these little cells as close as possible together. It is then easily conceivable that the symmetrical tensions of the adjacent films ... should suffice to bring the precise configuration which the comb actually presents.’ The laws of thermodynamics would cause the wax to achieve the optimal, energy-minimizing configuration, with no need for complex building behaviour finely honed by natural selection.



According to Oldroyd and Pratt:



> Thompson’s hypothesis is appealingly simple, but it does not seem consistent with currently available data on actual bees



> Detailed observation of building bees makes clear that they actively bite, form, and plane the developing cells, rather than forming a passive matrix around which wax flows. Further, hexagonal cells are also typical of social wasps like Polistes and Vespula. These cells are constructed from macerated plant material, a substance much less likely to have the thermoplastic properties of warm wax. These facts suggest that comb construction depends on context-dependent rules that the bees (and wasps) follow depending on the kind of cells required



> Cells under construction by Apis mellifera, show the highly regular array of hexagonal cells with rhomboid bases.



Oldroyd, B. P., & Pratt, S. C. (2015). Comb Architecture of the Eusocial Bees Arises from Simple Rules Used During Cell Building. Advances in Insect Physiology.



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