Do the protocadherins Fat and Dachsous link up to determine both planar cell polarity and the dimensions of organs?

Most, perhaps all cells in epithelial sheets are polarized in the plane of the sheet. This type of polarity, referred to as planar cell polarity (PCP), can be expressed in the orientation of cilia and stereocilia, in oriented outgrowths such as hairs, in the plane of cell division, in directed cell movement and possibly in the orientation of axon extension. Another popular area in current research is growth: there is an attempt to find systems that fix the shape and size of organs. Although both polarity and growth are subject to overall control by morphogen gradients, the mechanisms of this control are almost completely unknown. Here we discuss recent evidence for a 'steepness hypothesis' that links these two apparently disconnected features of animal development.

[1]  Peter A. Lawrence,et al.  Polarity and Patterns in the Postembryonic Development of Insects , 1970 .

[2]  P. Lawrence,et al.  Developmental Compartments and Planar Polarity in Drosophila , 2002, Current Biology.

[3]  J. Huxley Problems of relative growth , 1932 .

[4]  G. Halder,et al.  Boundaries of Dachsous Cadherin activity modulate the Hippo signaling pathway to induce cell proliferation , 2008, Proceedings of the National Academy of Sciences.

[5]  P J Bryant,et al.  Mutations at the fat locus interfere with cell proliferation control and epithelial morphogenesis in Drosophila. , 1988, Developmental biology.

[6]  Last hideout of the unknown? , 2004, Nature.

[7]  B. Edgar,et al.  Filling out the Hippo pathway , 2007, Nature Reviews Molecular Cell Biology.

[8]  K. Irvine,et al.  Four-jointed Is a Golgi Kinase That Phosphorylates a Subset of Cadherin Domains , 2008, Science.

[9]  P. Lawrence,et al.  Hedgehog acts by distinct gradient and signal relay mechanisms to organise cell type and cell polarity in the Drosophila abdomen. , 1997, Development.

[10]  P. Lawrence,et al.  Planar cell polarity: one or two pathways? , 2007, Nature Reviews Genetics.

[11]  Dragana Rogulja,et al.  Regulation of Cell Proliferation by a Morphogen Gradient , 2005, Cell.

[12]  R. Rosenfeld Nature , 2009, Otolaryngology--head and neck surgery : official journal of American Academy of Otolaryngology-Head and Neck Surgery.

[13]  F. Crick,et al.  A gradient of positional information in an insect, Rhodnius. , 1972, Journal of cell science.

[14]  P. Lawrence,et al.  Measuring dimensions: the regulation of size and shape. , 2000, Development.

[15]  Dragana Rogulja,et al.  Morphogen control of wing growth through the Fat signaling pathway. , 2008, Developmental cell.

[16]  A. Mccarthy Development , 1996, Current Opinion in Neurobiology.

[17]  G. Struhl,et al.  Recruitment of cells into the Drosophila wing primordium by a feed-forward circuit of vestigial autoregulation , 2007, Development.

[18]  Jennifer A Zallen,et al.  Planar Polarity and Tissue Morphogenesis , 2007, Cell.

[19]  G. Struhl,et al.  Control of Drosophila wing growth by the vestigial quadrant enhancer , 2007, Development.

[20]  N. Tapon,et al.  The Tumor-Suppressor Gene fat Controls Tissue Growth Upstream of Expanded in the Hippo Signaling Pathway , 2006, Current Biology.

[21]  David Strutt,et al.  Cleavage and secretion is not required for Four-jointed function in Drosophila patterning , 2004, Development.

[22]  K. Harvey,et al.  Fat Cadherin Modulates Organ Size in Drosophila via the Salvador/Warts/Hippo Signaling Pathway , 2006, Current Biology.

[23]  M. Simon Planar cell polarity in the Drosophila eye is directed by graded Four-jointed and Dachsous expression , 2004, Development.

[24]  Yaopan Mao,et al.  Dachs: an unconventional myosin that functions downstream of Fat to regulate growth, affinity and gene expression in Drosophila , 2006, Development.

[25]  G. Struhl,et al.  Sequential organizing activities of engrailed, hedgehog and decapentaplegic in the Drosophila wing. , 1995, Development.

[26]  R. Fehon,et al.  Delineation of a Fat tumor suppressor pathway , 2006, Nature Genetics.

[27]  K. Irvine,et al.  The Fat and Warts signaling pathways: new insights into their regulation, mechanism and conservation , 2008, Development.

[28]  C. Pantin Problems of Relative Growth , 1932, Nature.

[29]  I. Hariharan,et al.  Regulation of imaginal disc growth by tumor-suppressor genes in Drosophila. , 2006, Annual review of genetics.

[30]  Peter A Lawrence,et al.  Two separate molecular systems, Dachsous/Fat and Starry night/Frizzled, act independently to confer planar cell polarity , 2006, Development.

[31]  Claud L. Brown,et al.  Growth and Form , 1971 .

[32]  Peter J. Bryant,et al.  The fat tumor suppressor gene in Drosophila encodes a novel member of the cadherin gene superfamily , 1991, Cell.

[33]  D'arcy W. Thompson,et al.  On Growth and Form , 1917, Nature.

[34]  David Strutt,et al.  Nonautonomous planar polarity patterning in Drosophila: dishevelled-independent functions of frizzled. , 2002, Developmental cell.

[35]  Jeffrey D. Axelrod,et al.  Regulation of Frizzled by Fat-like Cadherins during Planar Polarity Signaling in the Drosophila Compound Eye , 2002, Cell.

[36]  Ryan S. Udan,et al.  The Fat Cadherin Acts through the Hippo Tumor-Suppressor Pathway to Regulate Tissue Size , 2006, Current Biology.

[37]  J. Nathans,et al.  Tissue/planar cell polarity in vertebrates: new insights and new questions , 2007, Development.

[38]  P. Lawrence Gradients in the Insect Segment: The Orientation of Hairs in the Milkweed Bug Oncopeltus Fasciatus , 1966 .

[39]  N. Perrimon,et al.  Multiple roles for four-jointed in planar polarity and limb patterning. , 2000, Developmental biology.