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Growth control in the Drosophila wing disk

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Abstract The regulation of size and shape is a fundamental requirement of biological development and has been a subject of scientific study for centuries, but we still lack an understanding of how organisms know when to stop growing. Imaginal wing disks of the fruit fly Drosophila melanogaster, which are precursors of the adult wings, are an archetypal tissue for studying growth control. The growth of the disks is dependent on many inter‐ and intra‐organ factors such as morphogens, mechanical forces, nutrient levels, and hormones that influence gene expression and cell growth. Extracellular signals are transduced into gene‐control signals via complex signal transduction networks, and since cells typically receive many different signals, a mechanism for integrating the signals is needed. Our understanding of the effect of morphogens on tissue‐level growth regulation via individual pathways has increased significantly in the last half century, but our understanding of how multiple biochemical and mechanical signals are integrated to determine whether or not a cell decides to divide is still rudimentary. Numerous fundamental questions are involved in understanding the decision‐making process, and here we review the major biochemical and mechanical pathways involved in disk development with a view toward providing a basis for beginning to understand how multiple signals can be integrated at the cell level, and how this translates into growth control at the level of the imaginal disk. This article is categorized under: Analytical and Computational Methods > Computational Methods Biological Mechanisms > Cell Signaling Models of Systems Properties and Processes > Cellular Models
(a) The wing disk—A: anterior, P: posterior, D: dorsal, V: ventral, AP and DV: anterior–posterior and dorsal‐ventral boundaries, DP: disk pouch. (b) Side view along B in (a). (Reprinted with permission from Widmann and Dahmann ())
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Other regulators of the Hippo pathway (Reprinted with permission from Zheng and Pan ())
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A 2D slice of the disk along its apicobasal axis (Meyer, Ikmi, & Gibson, ), yellow arrows are cells that are dividing. Note the large aspect ratio and curvature of the disk
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An apical section of a cell in a tissue, showing the E‐Cad junctions (green), the cortex, Arm (red), myosin (blue) and the apical CSK
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Images of a wing disk in which the apical cell outlines of the disk proper are marked by GFP fused to E‐Cadherin. In (a) the large muscle fiber between the wing, leg, and haltere disks is exerts a substantial force on the disk at (**). (b) shows the same wing disk after dissection, which relaxes the external force and leads to a significant change in cell shape. The scale bar is 50 mm. From (Nienhaus et al., )
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Spatial gene expression profiles for genes involved in BMP signaling. Modified from (Raftery & Umulis, )
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A cross‐section of a disk showing the transport processes that affect the morphogen distribution (Othmer, Painter, Umulis, & Xue, ). ZA are aka AJ
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The Dpp signaling pathway. The receptor is a complex of two Tkv and two Punt molecules
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The cell‐based Hippo pathway. (Reprinted with permission from Gou, Lin, and Othmer ()). See also Figure
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The Hh, Dpp, and Wg pathways. Arm: armadillo, Ban: bantam, Brk: brinker, Nkd: naked, Dad (Mad): daughters (mothers) against Dpp, Hpo: Hippo, Dfz: Drosophila frizzeled, Dlp: dally‐like protein, Gro: groucho, Hth: homothorax, Dsh: disheveled, Mtv: master of thickveins, Tkv: thickveins, Omb: optomotorblind, Pan: pangolin, Sd: scalloped, Smo: smoothened, Ubx: ultrabithorax. Assembled from (Crickmore & Mann, ; Fujise et al., ; Funakoshi, Minami, & Tabata, ; Held, ; Rodríguez, ; Sopko & McNeill, ; Zeng, Rahnama, Wang, Lee, & Verheyen, ). See also Table
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Models of Systems Properties and Processes > Cellular Models
Biological Mechanisms > Cell Signaling

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