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WIREs Syst Biol Med
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Branch formation during organ development

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Abstract Invertebrates and vertebrates use branching morphogenesis to build epithelial trees to maximize the surface area of organs within a given volume. Several molecular regulators of branching have recently been discovered, a number of which are conserved across different organs and species. Signals that control branching at the cellular and tissue levels are also starting to emerge, and are rapidly unveiling the physical nature of branch development. Here we discuss the molecular, cellular, and physical processes that govern branch formation, and highlight the major outstanding questions in the field. Copyright © 2010 John Wiley & Sons, Inc. This article is categorized under: Developmental Biology > Developmental Processes in Health and Disease

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Mechanical control of branching. The high mechanical fields at the tips enhance branching by physically propelling the epithelium forward and by activating proteases and signaling pathways that enhance cellular motility and invasiveness.

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Branch formation by viscous fingering. (a) The pressure in the epithelium is initially uniform, whereas that of the mesenchyme decreases away from the epithelium. (b) A small bulge in the epithelium protrudes and encounters a sharper pressure drop, which drives further protrusion. (c) As the bud grows, it displaces the mesenchyme toward the stalk regions, reducing the pressure drop there and lowering the driving force for motion.

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Branch formation by chemoattraction. (a) Fibroblast growth factor (FGF) source (orange) guides branch extension by enhancing motility of tip cells. FGF induces expression of Sonic hedgehog (Shh) (blue), which in turn suppresses and splits the FGF source. The split source of FGF gives rise to branch bifurcation. (b) Cells expressing high levels of Ret (green), persistently migrate toward the source of glial cell line‐derived neutrophic factor (GDNF) (purple), forming a patch of high Ret activity, which ultimately forms the ureteric bud.

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Organs constructed by branch formation. (a) Drosophila trachea, (b) chicken lung and (c) mouse kidney (Reprinted with permission from Ref 1. Copyright 2009 Elsevier), and (d) mouse salivary gland (Reprinted with permission from Ref 2. Copyright 2006 Macmillan Publishers Ltd.). Scale bars, 100 µm.

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