Guidebook to the Homeobox Genes. 1994.
Denis Duboule (ed.)
Oxford University Press, Oxford. ISBN 0-19-859939-0 (hardcover), 0-19-859940-4 (paperback).

This book also contains a chapter on the classification of homeobox genes written by myself that should give novices a good entry point into understanding all the different types of homeobox genes. (A Comprehensive Classification of Homeobox Genes, Bürglin, T.R. (1994) in Guidebook to the Homeobox Genes. Duboule, D. (ed.) Oxford University Press, Oxford pp 25-71). At present it is out of print.

Although a lot of information is out of date at this point, it still has some useful information not found in later papers.
Here on this page some errors and changes are pointed out.

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  • page 66: NvHbox2.8 has been assigned as HoxA2 by Swissprot. With blast searches it does come out closer to HoxA2 genes. But it should be linked to NvHbox2.7. What is it really?
  • page 48: Fig 10: h HOXB3 is mislabelled; should be h HOXD3
  • page 17: Pax/Prd family: Pax-6 is small-eyes, and Pax-3 is splotch (was switched)
  • page 89: Athb-1 is HAT5 (page 134), reference is Ruberti, not Roberti as listed in references.
  • page 122: EST01828 is human OTX2
  • page 43: m HoxD3 sequence is SKRV... instead of SKSV...
  • page 48: the human HB9 sequence was apparently incorrect, the latest database entry does not have the gap of 2 amino acids (X56537), thus Figure 10 can be ignored.

  • Note: one large subgroup of atypical homeodomains is the TALE group that was defined later.


  • The Evolution of Homeobox genes. Bürglin, T.R. (1995) in 'Biodiversity and Evolution', Arai, R., Kato, M., Doi, Y. (eds.), The National Science Museum Foundation, Tokyo, pp. 291-336, ISBN 4-9900433-0-8, The National Science Museum Foundation, 7-20 Ueno Park, Taito-ku, Tokyo 110, Japan.

    Highlights from that paper:

  • Comparative tree of classes figure.

  • Columns on the right mark in what groups of animals classes have been found: d: Drosophila, v: vertebrates, c: C. elegans, o: others, i.e. leech, sea urchin, molluscs, planaria etc.
  • Classes with conserved sequence domains/motifs figure.
  • New classes: ro (rough), so (sine oculis), Mox, Lb (lady-bird).
  • TCL class renamed into Tlx class.
  • New atypical class: pros (prospero); with conserved domain carboxy-terminal to the homeodomain.
  • New domain/motif: so class has a long conserved domain, the Six/so domain, upstream of the homeodomain.
  • All atypical homeobox genes that have 3 extra amino acids between helix 1 and helix 2 have been grouped together (This group is now known as the  TALE superclass of homeobox genes). They have been classified into 4 groups: the established PBC class, the new M-ATYP class for the fungal mating type genes, the Kn (Knotted) class for this group of plant genes, and the HAC-ATYP class, a novel class that is conserved between animals and plants. A figure of these sequences shows the conservation between helix 1 and helix 2.
  •           Note about the classification: Subsequently names and classification were revised for TALE homeobox genes (e.g., HAC-ATYP split into BEL and MEIS).

              Notes and corrections for that paper:
  • In Figure 1B, in the first column the 'K' is missing in the light grey area.
  • The Kn class is now called the KNOX class (see below). HAC-ATYP is abandoned due to a revised nomenclature (see TALE paper below).