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Key publications

  • Helen Weavers, Silvia Prieto-Sánchez, Ferdinand Grawe, Amparo Garcia-López, Ruben Artero, Michaela Wilsch-Braeuninger, Mar Ruiz-Gómez, Helen Skaer & Barry Denholm (2009) The insect nephrocyte is a podocyte-like cell with a filtration slit diaphragm. Nature 457:322-326
  • Denholm, B. et al (2013) The tiptop/teashirt genes regulate cell differentiation and renal physiology in Drosophila Development 140:1100-1110; doi:10.1242/dev.088989   
  • Weavers, H. and Skaer, H. (2013) Tip cells act as dynamic cellular anchors in the morphogenesis of looped renal tubules in Drosophila. Developmental Cell 27: 331–344.
  • Saxena, A, Denholm, B, Bunt, S, Bischoff, M, VijayRaghavan, K and Skaer, H  (2014) Epidermal growth factor signalling controls Myosin II planar polarity to orchestrate convergent-extension movements during Drosophila tubulogenesis PLOS Biology in press
  • Bunt, S. et al (2010) Haemocyte-secreted Type IV Collagen enhances BMP signalling to guide renal tubule morphogenesis in Drosophila Developmental Cell 19: 296-306. doi: 10.1016/j.devcel.2010.07.019


Other selected publications

  • Skaer, H. (1989) Cell division in the development of the Malpighian tubules of Drosophila melanogaster is regulated by single, specialised cells.  Nature 342, 566-569.
  • Skaer, H., Harrison, J.B. & Maddrell, S.H.P. (1990) Physiological and structural maturation of a polarised epithelium - the Malpighian tubules of a blood-sucking insect, Rhodnius prolixus.  J. Cell Sci. 96, 537-548
  • Skaer, H. (1992) Cell proliferation and rearrangement in the development of the Malpighian tubules of the Hemipteran, Rhodnius  prolixus. Dev. Biol. 150, 372-380.
  • Broadie, K., Skaer, H. & Bate, M. (1992) Whole-embryo culture of Drosophila: development of embryonic tissues in vitro. Roux' Arch. Dev. Biol. 201, 364-375
  • Skaer, H, and Martinez Arias, A. (1992) The wingless product is required for cell proliferation in the Malpighian tubule anlage of Drosophila melanogaster Development 116, 745-754
  • Baumann, P and Skaer, H. (1993) The Drosophila EGF receptor  homolgue (DER) is required for Malpighian tubule development.  Development Suppl. 65 - 75.
  • Hoch, M., Broadie, K., Jäckle, H. and Skaer, H. (1994) Sequential fates in a single cell are established by the neurogenic cascade in the Malpighian tubules of Drosophila.  Development 120, 3439 - 3450.
  • Grawe, F., Wodarz, A., Lee, W., Knust, E. and Skaer, H. (1996) The Drosophila genes crumbs and stardust are involved in the biogenesis of adherens junctions. Development. 122, 951 - 959.
  • Skaer, H. (1996). Cell proliferation and development of the Malpighian tubules in Drosophila melanogaster. Experimental Nephrology  4, 119 – 126
  • Denholm, B. Brown, S., Ray, R., Ruiz-Gómez, M., Skaer, H. and Castelli-Gair Hombría, J. (2005) crossveinless-c is a RhoGAP required for actin reorganisation during morphogenesis. Development  132: 2389-2400.
  • Simões S, Denholm B, Azevedo D, Sotillos S, Martin P, Skaer H, Hombría JC, Jacinto A. (2006) Apical Rho-GTPase activation required for epithelial invagination is controlled by opposing localization of GEF and GAP regulators to the apical and basal membrane domains. Development 133: 4257-67.
  • Hatton-Ellis, E., Ainsworth, C., Sushama, Y., Wan, S., VijayRaghavan, K. and Skaer, H. (2007) The genetic regulation of patterned tubular branching in Drosophila. PNAS 104;169-174
  • Xavier Caubit, Claire M. Lye, Elise Martin, Nathalie Coré, David A. Long, Christine Vola, Dagan Jenkins, Alistair N. Garratt, Helen Skaer, Adrian S. Woolf, and Laurent Fasano (2008) Teashirt 3 is necessary for ureteral smooth muscle differentiation downstream of SHH and BMP4 Development 135: 3301-3310.
  • Campbell, K., Knust, E. and Skaer, H. (2009) Crumbs stabilises epithelial polarity during tissue remodelling. J, Cell Sci. 122:2604-2612.
  • Campbell, K., Casanova, J. and Skaer, H. (2010) Mesenchymal-to-epithelial transition of intercalating cells in Drosophila renal tubules depends on polarity cues from epithelial neighbours. Mechanisms of Development 127: 345-357.
  • Johnson, K., Knust, E. and Skaer, H. (1999) bloated tubules (blot) encodes a Drosophila member of the neurotransmitter transporter family required for organisation of the apical cytocortex. Dev. Biol. 212: 440-454.
  • Wan, S., Cato, A-M. and Skaer, H. (2000) Multiple signalling pathways establish cell fate and number in Drosophila Malpighian tubules. Dev. Biol. 217: 153-165.
  • Ainsworth, C., Wan, S. &  Skaer, H. (2000) Coordinating cell fate and morphogenesis in Drosophila renal tubules. Phil Trans Roy Soc (series B) 355: 931 – 937.
  • Sudarsan, V., Anant, S., Guptan, P., VijayRaghavan, K. and Skaer, H. (2001) Myoblast diversification and ectodermal signaling in Drosophila. Developmental Cell. 1: 829 – 840.
  • Sudarsan, V., Pasalodos-Sanchez, S., Wan, S., Gampel, A. and Skaer, H. (2002)  A genetic hierarchy establishes mitogenic signalling and mitotic competence in the renal tubules of Drosophila. Development. 129: 935-944.
  • Denholm, B., Sudarsan, V., Pasalodos Sanchez, S., Artero, R., Lawrence, P, Maddrell, S., Baylies, M. and Skaer, H. (2003) Dual origin of the renal tubules in Drosophila: mesodermal cells integrate and polarise to establish secretory function.  Curr. Biol. 13: 1052-1057
  • Bunt, S., Hooley, C., Hu, N., Scahill, C., Weavers, H. and Skaer, H. (2010) Haemocyte-secreted Type IV Collagen enhances BMP signalling to guide renal tubule morphogenesis in Drosophila Developmental Cell 19: 296-306.
  • Bunt, S., Denholm, D. and Skaer, H. (2011) Characterisation of the Drosophila procollagen lysyl hydroxylase, D-Plod. Gene Expression Patterns 11: 72-78


  • Beyenbach, K. W., Skaer, H. and Dow, J.A.T. (2009) Developmental, Molecular and Transport Biology of Malpighian tubules. Annu. Rev. Entomol. 2010. 55:351–74.
  • Denholm, B. and Skaer, H. (2009) Bringing together components of the fly renal system. Current Opinion in Genetics & Development 2009, 19:526–532.
  • Weavers, H and Skaer, H (2014) Tip cells: master regulators of tubulogenesis? Seminars in Cell and Developmental Biology 31: 91-99