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WIREs Comput Mol Sci
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Eukaryotic cell dynamics from crawlers to swimmers

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Movement requires force transmission to the environment, and motile cells are robustly, though not elegantly, designed nanomachines that often can cope with a variety of environmental conditions by altering the mode of force transmission used.a As with humans, the available modes range from momentary attachment to a substrate when crawling, to shape deformations when swimming, and at the cellular level this involves sensing the mechanical properties of the environment and altering the mode appropriately. While many types of cells can adapt their mode of movement to their microenvironment (ME), our understanding of how they detect, transduce and process information from the ME to determine the optimal mode is still rudimentary. The shape and integrity of a cell is determined by its cytoskeleton (CSK), and thus the shape changes that may be required to move involve controlled remodeling of the CSK. Motion in vivo is often in response to extracellular signals, which requires the ability to detect such signals and transduce them into the shape changes and force generation needed for movement. Thus the nanomachine is complex, and while much is known about individual components involved in movement, an integrated understanding of motility in even simple cells such as bacteria is not at hand. In this review we discuss recent advances in our understanding of cell motility and some of the problems remaining to be solved. This article is categorized under Structure and Mechanism > Computational Materials Science Structure and Mechanism > Computational Biochemistry and Biophysics
(a) Four time points of a swimming Dictyostelium discoideum cell showing the axial propagation of protrusions along the length. (Reprinted with permission from Ref . Copyright 2010 National Academy of Science) (b) Three time frames from a computational model of a two‐dimensional swimmer using symmetric protrusions. (c) The mean velocity of two swimmers as a function of the protrusion height. (b) and (c) (Reprinted with permission from Ref . Copyright 2015 Springer)
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The push‐me‐pull‐you swimmer
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The ratio of Ras* under uniform stimulation to its basal level as a function of the stimulus level and time. (Reprinted with permission from Ref . Copyright 2016 Public Library of Science)
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The levels of activated Ras* at the front and back in a static cAMP gradient as a function of time measured experimentally (a) (Reprinted with permission from Ref . Copyright 2013 Company of Biologists) and the model prediction (b). (c) The average Ras* in the front and rear halves in response to a passing triangle wave. (Reprinted with permission from Ref . Copyright 2016 Public Library of Science)
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A schematic of the major processes in the model (top), and the major steps in the network (bottom). (Reprinted with permission from Ref . Copyright 2016 Public Library of Science)
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(a) A cross‐section of the waves. (Reprinted with permission from Ref . Copyright 2010 Nature Publishing Group) A schematic of the network structure and molecular interactions in the model. Two snapshots in time of an actin wave initiated at x = 2.5, showing the network density (color) as a function of space (x‐axis) and network height (z‐axis). ((b) and (c) Reprinted with permission from Ref . Copyright 2013 Public Library of Science)
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(a) Some of the major components of cAMP signal transduction in Dictyostelium discoideum (Dd). CAR1: the cAMP receptor, Gαβγ: a G‐protein involved in the transduction of the extracellular signal, Ras: a small G‐protein, PIP2, and PIP3; components of the membrane that can be interconverted via phosphorylation and dephosphorylation, IP3 and DAG: products that result from the degradation of PIP2, Ca2: calcium, GC: guanylate cyclase—the enzyme that produces cyclic GMP (cGMP), AC: adenylate cyclase—the enzyme that produces cAMP, Rac1: a small G‐rotein, Myosin: a motor protein involved in contraction of the actin network. (b) The PIP2–PIP3 trio. Activated Ras activates PI3K, which phosphorylates PIP2. PIP3 provides a binding site for cytosolic PI3K, thereby creating a positive feedback loop through PI3K. Similarly, PIP2 provides a binding site for PTEN, which dephosphorylates PIP3. PIP3 levels are controlled in part by PTEN and SHIP, which dephosphorylate PIP3 at different sites. (c) The skeletal network downstream of Ras that determines the balance between dendritic network formation and myo‐II assembly in Dd
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(a) The normal versus the in‐plane force in Dictyostelium discoideum (Dd). The inset shows the labeling. (b) The cyclic variation of the forces and the speed. (c) and (d) The tangential and normal forces in Dd. (a) and (b) (Reprinted with permission from Ref . Copyright 2010 American Physical Society) (c) and (d) (Reprinted with permission from Ref . Copyright 2015 Cell Press)
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How the shape of a keratocyte depends on the subtrate stiffness. (Reprinted with permission from Ref . Copyright 2016 Nature Publishing Group)
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The molecular clutch model. (Reprinted with permission from Ref . Copyright 2008 American Association for the Advancement of Science)
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A schematic of a mesenchymal cell, showing the various substructures and protrusions. (Reprinted with permission from Ref . Copyright 2014 American Physiological Society)
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(a) Blebbing on a melanoma cell: myosin (green) localizes under the blebbing membrane (red). (b) The actin cortex of a Dictyostelium discoideum cell migrating to the lower right. Arrowheads indicate the successive blebs and arcs of the actin cortex. (Reprinted with permission from Ref . Copyright 2008 Nature Publishing Group)
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Two examples of mesenchymal motion. Top: A fibroblast in a three‐dmensional collagen matrix. (Reprinted with permission from Ref . Copyright 2012 Company of Biologists) Bottom: A fish‐scale keratocyte. Actin (green) myosin‐II (red) and focal adhesions (blue). From www.hhmi.org/scientists/julie‐theriot
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Top: The type A1 morphology and the cortical flow rates observed in zebrafish progenitor cells. (Reprinted with permission from Ref . Copyright 2015 Cell Press) Bottom: A schematic of the cortical and interior flow
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A summary of the different modes of movement in different environments and under different substrate properties. (Reprinted with permission from Ref . Copyright 2016 Elsevier Ltd)
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Structure and Mechanism > Computational Biochemistry and Biophysics
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