, Volume 100, Issue 1, pp 45–49 | Cite as

Hexagonal comb cells of honeybees are not produced via a liquid equilibrium process

  • Daniel Bauer
  • Kaspar BienefeldEmail author
Original Paper


The nests of European honeybees (Apis mellifera) are organised into wax combs that contain many cells with a hexagonal structure. Many previous studies on comb-building behaviour have been made in order to understand how bees produce this geometrical structure; however, it still remains a mystery. Direct construction of hexagons by bees was suggested previously, while a recent hypothesis postulated the self-organised construction of hexagonal comb cell arrays; however, infrared and thermographic video observations of comb building in the present study failed to support the self-organisation hypothesis because bees were shown to be engaged in direct construction. Bees used their antennae, mandibles and legs in a regular sequence to manipulate the wax, while some bees supported their work by actively warming the wax. During the construction of hexagonal cells, the wax temperature was between 33.6 and 37.6 °C. This is well below 40 °C, i.e. the temperature at which wax is assumed to exist in the liquid equilibrium that is essential for self-organised building.


Cell geometry Comb building Honeybee comb Liquid equilibrium hypothesis 



We thank Uwe Gerber for the support with constructing the observation unit. We also thank two anonymous referees for their valuable comments. This work was supported by funds from the regional governments of Brandenburg, Berlin, Saxony, Saxony-Anhalt and Thuringia. Part of the equipment used was financed by the European Union (EFRE–Application No. 80137041).

Supplementary material

114_2012_992_Fig4_ESM.jpg (7 kb)
Online Resource 1

Assembly of the observation hive. The hive was open on one side. A digital infrared camera (a) and a thermographic camera (b) projected directly into the box facing the comb without any foil in between. The box was closed and thermally insulated with styrofoam. Thermo elements (c) and a sensor (d) were used to maintain the temperature manually at ca 30 °C. A glass plate was placed between the cameras to prevent heat transmission. An infrared light source (e) was placed at the bottom (JPEG 6 kb)

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High resolution image (TIFF 1769 kb)
Online Resource 2

Cross section of a comb containing building bees. In this infrared video, the building technique of single bees is shown inside and above the cells (AVI 22777 kb)

Online Resource 3

Thermographic view of comb building. This video shows the comb building behaviour in a thermographic view. The colour scale provides information about the temperature. A cell was excised in each of windows 1 and 2, and builders performed repair work. The area with warm bees below is the comb edge (AVI 11752 kb)

114_2012_992_Fig5_ESM.jpg (6 kb)
Online Resource 4

Hexagonal comb cells built by the social wasp Dolichovespula saxonica. Wasps produce hexagonal cells using cellulose as the building material (JPEG 5 kb)

114_2012_992_MOESM4_ESM.tif (393 kb)
High resolution image (TIFF 392 kb)


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Copyright information

© Springer-Verlag Berlin Heidelberg 2012

Authors and Affiliations

  1. 1.Institute for Bee Research Hohen NeuendorfHohen NeuendorfGermany

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