How Cells Integrate Complex Stimuli: The Effect of Feedback from Phosphoinositides and Cell Shape on Cell Polarization and Motility
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{"title"=>"How cells integrate complex stimuli: The effect of feedback from phosphoinositides and cell shape on cell polarization and motility", "type"=>"journal", "authors"=>[{"first_name"=>"Athanasius F M", "last_name"=>"Marée", "scopus_author_id"=>"6602862491"}, {"first_name"=>"Verônica A.", "last_name"=>"Grieneisen", "scopus_author_id"=>"14060094700"}, {"first_name"=>"Leah", "last_name"=>"Edelstein-Keshet", "scopus_author_id"=>"6701669583"}], "year"=>2012, "source"=>"PLoS Computational Biology", "identifiers"=>{"pmid"=>"22396633", "sgr"=>"84861146594", "doi"=>"10.1371/journal.pcbi.1002402", "scopus"=>"2-s2.0-84861146594", "pui"=>"364830942", "isbn"=>"1553-734x", "issn"=>"1553734X"}, "id"=>"e0fbd668-bf52-3704-8563-279ed0b196ae", "abstract"=>"To regulate shape changes, motility and chemotaxis in eukaryotic cells, signal transduction pathways channel extracellular stimuli to the reorganization of the actin cytoskeleton. The complexity of such networks makes it difficult to understand the roles of individual components, let alone their interactions and multiple feedbacks within a given layer and between layers of signalling. Even more challenging is the question of if and how the shape of the cell affects and is affected by this internal spatiotemporal reorganization. Here we build on our previous 2D cell motility model where signalling from the Rho family GTPases (Cdc42, Rac, and Rho) was shown to organize the cell polarization, actin reorganization, shape change, and motility in simple gradients. We extend this work in two ways: First, we investigate the effects of the feedback between the phosphoinositides (PIs) PIP₂, PIP₃ and Rho family GTPases. We show how that feedback increases heights and breadths of zones of Cdc42 activity, facilitating global communication between competing cell \"fronts\". This hastens the commitment to a single lamellipodium initiated in response to multiple, complex, or rapidly changing stimuli. Second, we show how cell shape feeds back on internal distribution of GTPases. Constraints on chemical isocline curvature imposed by boundary conditions results in the fact that dynamic cell shape leads to faster biochemical redistribution when the cell is repolarized. Cells with frozen cytoskeleton, and static shapes, consequently respond more slowly to reorienting stimuli than cells with dynamic shape changes, the degree of the shape-induced effects being proportional to the extent of cell deformation. We explain these concepts in the context of several in silico experiments using our 2D computational cell model.", "link"=>"http://www.mendeley.com/research/cells-integrate-complex-stimuli-effect-feedback-phosphoinositides-cell-shape-cell-polarization-motil", "reader_count"=>128, "reader_count_by_academic_status"=>{"Unspecified"=>3, "Professor > Associate Professor"=>12, "Librarian"=>1, "Researcher"=>37, "Student > Doctoral Student"=>5, "Student > Ph. D. Student"=>47, "Student > Postgraduate"=>3, "Student > Master"=>9, "Other"=>1, "Student > Bachelor"=>3, "Lecturer"=>1, "Lecturer > Senior Lecturer"=>2, "Professor"=>4}, "reader_count_by_user_role"=>{"Unspecified"=>3, "Professor > Associate Professor"=>12, "Librarian"=>1, "Researcher"=>37, "Student > Doctoral Student"=>5, "Student > Ph. D. Student"=>47, "Student > Postgraduate"=>3, "Student > Master"=>9, "Other"=>1, "Student > Bachelor"=>3, "Lecturer"=>1, "Lecturer > Senior Lecturer"=>2, "Professor"=>4}, "reader_count_by_subject_area"=>{"Engineering"=>12, "Unspecified"=>3, "Biochemistry, Genetics and Molecular Biology"=>12, "Mathematics"=>11, "Agricultural and Biological Sciences"=>57, "Arts and Humanities"=>2, "Neuroscience"=>1, "Physics and Astronomy"=>22, "Chemical Engineering"=>2, "Psychology"=>2, "Social Sciences"=>1, "Computer Science"=>3}, "reader_count_by_subdiscipline"=>{"Engineering"=>{"Engineering"=>12}, "Neuroscience"=>{"Neuroscience"=>1}, "Social Sciences"=>{"Social Sciences"=>1}, "Physics and Astronomy"=>{"Physics and Astronomy"=>22}, "Psychology"=>{"Psychology"=>2}, "Agricultural and Biological Sciences"=>{"Agricultural and Biological Sciences"=>57}, "Computer Science"=>{"Computer Science"=>3}, "Biochemistry, Genetics and Molecular Biology"=>{"Biochemistry, Genetics and Molecular Biology"=>12}, "Mathematics"=>{"Mathematics"=>11}, "Unspecified"=>{"Unspecified"=>3}, "Chemical Engineering"=>{"Chemical Engineering"=>2}, "Arts and Humanities"=>{"Arts and Humanities"=>2}}, "reader_count_by_country"=>{"Canada"=>1, "Sweden"=>1, "Argentina"=>1, "United States"=>2, "Japan"=>1, "Brazil"=>1, "United Kingdom"=>2, "France"=>1, "Portugal"=>3, "Germany"=>2, "Spain"=>1, "India"=>1}, "group_count"=>4}

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Figshare

  • {"files"=>["https://ndownloader.figshare.com/files/346595", "https://ndownloader.figshare.com/files/346690", "https://ndownloader.figshare.com/files/346793", "https://ndownloader.figshare.com/files/346900", "https://ndownloader.figshare.com/files/346995", "https://ndownloader.figshare.com/files/347085", "https://ndownloader.figshare.com/files/347238", "https://ndownloader.figshare.com/files/347330", "https://ndownloader.figshare.com/files/347454", "https://ndownloader.figshare.com/files/347559"], "description"=>"<div><p>To regulate shape changes, motility and chemotaxis in eukaryotic cells, signal transduction pathways channel extracellular stimuli to the reorganization of the actin cytoskeleton. The complexity of such networks makes it difficult to understand the roles of individual components, let alone their interactions and multiple feedbacks within a given layer and between layers of signalling. Even more challenging is the question of if and how the shape of the cell affects and is affected by this internal spatiotemporal reorganization. Here we build on our previous 2D cell motility model where signalling from the Rho family GTPases (Cdc42, Rac, and Rho) was shown to organize the cell polarization, actin reorganization, shape change, and motility in simple gradients. We extend this work in two ways: First, we investigate the effects of the feedback between the phosphoinositides (PIs) , and Rho family GTPases. We show how that feedback increases heights and breadths of zones of Cdc42 activity, facilitating global communication between competing cell “fronts”. This hastens the commitment to a single lamellipodium initiated in response to multiple, complex, or rapidly changing stimuli. Second, we show how cell shape feeds back on internal distribution of GTPases. Constraints on chemical isocline curvature imposed by boundary conditions results in the fact that dynamic cell shape leads to faster biochemical redistribution when the cell is repolarized. Cells with frozen cytoskeleton, and static shapes, consequently respond more slowly to reorienting stimuli than cells with dynamic shape changes, the degree of the shape-induced effects being proportional to the extent of cell deformation. We explain these concepts in the context of several <em>in silico</em> experiments using our 2D computational cell model.</p> </div>", "links"=>[], "tags"=>["cells", "phosphoinositides", "polarization", "motility"], "article_id"=>128522, "categories"=>["Biological Sciences", "Biophysics"], "users"=>["Athanasius F. M. Marée", "Verônica A. Grieneisen", "Leah Edelstein-Keshet"], "doi"=>["https://dx.doi.org/10.1371/journal.pcbi.1002402.s001", "https://dx.doi.org/10.1371/journal.pcbi.1002402.s002", "https://dx.doi.org/10.1371/journal.pcbi.1002402.s003", "https://dx.doi.org/10.1371/journal.pcbi.1002402.s004", "https://dx.doi.org/10.1371/journal.pcbi.1002402.s005", "https://dx.doi.org/10.1371/journal.pcbi.1002402.s006", "https://dx.doi.org/10.1371/journal.pcbi.1002402.s007", "https://dx.doi.org/10.1371/journal.pcbi.1002402.s008", "https://dx.doi.org/10.1371/journal.pcbi.1002402.s009", "https://dx.doi.org/10.1371/journal.pcbi.1002402.s010"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/How_Cells_Integrate_Complex_Stimuli_The_Effect_of_Feedback_from_Phosphoinositides_and_Cell_Shape_on_Cell_Polarization_and_Motility/128522", "title"=>"How Cells Integrate Complex Stimuli: The Effect of Feedback from Phosphoinositides and Cell Shape on Cell Polarization and Motility", "pos_in_sequence"=>0, "defined_type"=>4, "published_date"=>"2012-03-01 02:22:02"}
  • {"files"=>["https://ndownloader.figshare.com/files/672558"], "description"=>"<p>The top row represents small GTPases, the middle row depicts phosphoinositides, and at the lowest level are the cytoskeletal components. Here we explore the effects of feedback from the PIs to the small GTPases, indicated by the red dashed lines. The parameter in the model represents the magnitude of the feedback ( means the feedback is absent, means it is essential for activating Cdc42 and Rac).</p>", "links"=>[], "tags"=>["pathways", "assumed"], "article_id"=>343052, "categories"=>["Biological Sciences", "Biophysics"], "users"=>["Athanasius F. M. Marée", "Verônica A. Grieneisen", "Leah Edelstein-Keshet"], "doi"=>["https://dx.doi.org/10.1371/journal.pcbi.1002402.g001"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Signalling_pathways_assumed_in_the_model_/343052", "title"=>"Signalling pathways assumed in the model.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2012-03-01 00:50:52"}
  • {"files"=>["https://ndownloader.figshare.com/files/672695"], "description"=>"<p>The cell is initiated at rest, stimulated for 10 sec with a 15% gradient in the Cdc42 basal activation rates, and simulated for a total of 8 minutes, after which the stimulus is removed. (See <a href=\"http://www.ploscompbiol.org/article/info:doi/10.1371/journal.pcbi.1002402#s4\" target=\"_blank\">Materials and Methods</a> for details.) The figure presents a snapshot of the distributions and profiles when the cell has reached an effective steady state, i.e. when the cell shape and profiles do not further change, except for small fluctuations due to the stochastic nature of the formalism. The following is shown in the panels from left to right. Top row: intracellular steady-state distributions of active Rho GTPases Cdc42, Rac, and Rho, for the polarized cell state, followed by a graph of the steady-state profiles along the front-back axis of the cell; Second row: inactive Rho GTPases, and corresponding profiles; Third row: phosphoinositides PI, PIP2, PIP3 and corresponding profiles. Fourth row: barbed end density, Arp2/3, F-actin density, filament orientation and barbed end orientation distributions. Fifth row: left graph shows the velocity over time of a polarized (black) and a resting cell (red). The rest state, which is the second possible steady state in this system, is stable against low-amplitude noise. The corresponding distributions of filament density, barbed end and actin filament orientation for the resting cell are shown at the right. Colour map: (box) Cytoskeleton orientation is encoded using a colour-wheel where each orientation is represented by a colour at the given angle along the circle (hue). The filament density ranges from black (no filaments) to maximal density at the greatest colour intensity; distribution of filament orientations varies in saturation from white (complete anisotropy) to the maximal colour intensity (total coherence). (lower colour bar) Scale used for relative concentration/density “heat maps” in this figure. Note that the steepness of the internal gradient of signalling chemicals is reflected in the tightness of the transition between hues. The “front-back” interface is here taken as the isocline shown in green (labelled “mean”). Green is used to represent the mean rest state concentration value for any of the PIs, small GTPases, and Arp2/3. Deviations from this mean are captured by the heat map, in which the percentage variations above or below the mean, are as indicated by the arrows along this bar. (See also <a href=\"http://www.ploscompbiol.org/article/info:doi/10.1371/journal.pcbi.1002402#pcbi.1002402.s001\" target=\"_blank\">Video S1</a>.)</p>", "links"=>[], "tags"=>["motility", "phenotype", "produced"], "article_id"=>343185, "categories"=>["Biological Sciences", "Biophysics"], "users"=>["Athanasius F. M. Marée", "Verônica A. Grieneisen", "Leah Edelstein-Keshet"], "doi"=>["https://dx.doi.org/10.1371/journal.pcbi.1002402.g002"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Basic_motility_phenotype_produced_by_the_8220_wildtype_8221_model_cell_/343185", "title"=>"Basic motility phenotype produced by the “wildtype” model cell.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2012-03-01 00:53:05"}
  • {"files"=>["https://ndownloader.figshare.com/files/672881"], "description"=>"<p>Shown are the Rac distributions in the 2D cell [(<b>a</b>) and <a href=\"http://www.ploscompbiol.org/article/info:doi/10.1371/journal.pcbi.1002402#pcbi.1002402.s001\" target=\"_blank\">Video S1</a>] and in a representative 1D cross-section along the cell diameter (<b>b,c</b>). Left panels: absent PI feedback (). Right panels: with PI feedback (). The sharp transition between high and low Rac activity is seen on the left (contours closely spaced), whereas PIs create a broader transition zone (right). Panels (b–c): schematics of differences in intracellular patterns due to maximal PI feedback to Rho proteins. (b): Inactive Rac (dot-dashed line) is nearly uniform for , but shows significant depletion close to the “front” for high . Decreasing the rate of diffusion of inactive Rac (red curves) has little effect on the profile when . In contrast, decreasing when leads to a lower peak of active Rac at the front. Panels (c): Communication of multiple peaks of active Rac is very slow in the case, and much more significant in the case . Hence, feedback from PIs helps to resolve conflicting cell “fronts”.</p>", "links"=>[], "tags"=>["pis", "gtpase"], "article_id"=>343375, "categories"=>["Biological Sciences", "Biophysics"], "users"=>["Athanasius F. M. Marée", "Verônica A. Grieneisen", "Leah Edelstein-Keshet"], "doi"=>["https://dx.doi.org/10.1371/journal.pcbi.1002402.g003"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Schematics_of_how_feedback_from_PIs_change_small_GTPase_profiles_/343375", "title"=>"Schematics of how feedback from PIs change small GTPase profiles.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2012-03-01 00:56:15"}
  • {"files"=>["https://ndownloader.figshare.com/files/672989"], "description"=>"<p>Shown are Cdc42 distribution profiles at indicated times using the colour scheme as in <a href=\"http://www.ploscompbiol.org/article/info:doi/10.1371/journal.pcbi.1002402#pcbi-1002402-g002\" target=\"_blank\">Fig. 2</a>. (<b>a</b>) (and <a href=\"http://www.ploscompbiol.org/article/info:doi/10.1371/journal.pcbi.1002402#pcbi.1002402.s002\" target=\"_blank\">Video S2</a>) Effect of cell shape on repolarization in the presence (, left) and absence (, right) of feedback from PIs to Rho GTPases. Cells with elliptical, static shape are initially polarized along their short axis using the standard, transient gradient protocol. Due to their shape alone (with no further stimulus or bias), there is a clear tendency for the cells to repolarize. The dynamics of shape-induced repolarization occurs much more rapidly when PI feedback is included. (<b>b</b>) Static circular cell in which the intracellular profiles have been modified into a highly curved profile. Over time, the curvature of the front-back interface flattens (with regions of higher curvature changing faster). (<b>c</b>) Local injection of Cdc42 appears to locally distort the intracellular interface, which then straightens again. Shape-sensitivity and robustness to local perturbations can be understood through the tendency of the reaction-diffusion system to minimize the front-back interface. The no-flux boundary conditions further assure that the all level curves (interfaces) maintain right-angles to the cell membrane. For the dynamics of (b,c), see also <a href=\"http://www.ploscompbiol.org/article/info:doi/10.1371/journal.pcbi.1002402#pcbi.1002402.s003\" target=\"_blank\">Video S3</a>.</p>", "links"=>[], "tags"=>["intracellular", "interface"], "article_id"=>343477, "categories"=>["Biological Sciences", "Biophysics"], "users"=>["Athanasius F. M. Marée", "Verônica A. Grieneisen", "Leah Edelstein-Keshet"], "doi"=>["https://dx.doi.org/10.1371/journal.pcbi.1002402.g004"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Effect_of_cell_shape_and_intracellular_8216_front_back_8217_interface_curvatures_/343477", "title"=>"Effect of cell shape and intracellular ‘front-back’ interface curvatures.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2012-03-01 00:57:57"}
  • {"files"=>["https://ndownloader.figshare.com/files/673071"], "description"=>"<p>(<b>a–b</b>) A comparison of Cdc42 repolarization in a cell whose shape is frozen (left columns in each sequence) with a control cell that has a dynamic shape (right columns in each sequence). (See also <a href=\"http://www.ploscompbiol.org/article/info:doi/10.1371/journal.pcbi.1002402#pcbi.1002402.s004\" target=\"_blank\">Video S4</a>.) In both cases, the cell is initially polarized by a transient gradient, then repolarized by either an orthogonal (a) or opposing (b) second gradient. Images on the left and right were taken at the same times after the second stimulus. Note that in (a), after 3 min there is a noticeable difference in the cell's polarity as seen from the angle of the front-back interface with the stimulus gradient; the difference is accentuated even further by 6 min. In (b), the static cell only partially repolarizes during the time span when the control cell has completely repolarized. The evolving cell shape spontaneously twists the intracellular interface which has to maintain its orthogonality to the cell edge. The increased curvature of this interface has a faster rate of flattening, driving the chemical dynamics to adjust more rapidly. As the chemistry also feeds back to protrusion/contraction and shape change, the two-way feedback resulting from cell-shape dynamics leads to a faster overall turning and aligning with the repolarization cue.</p>", "links"=>[], "tags"=>["intracellular"], "article_id"=>343564, "categories"=>["Biological Sciences", "Biophysics"], "users"=>["Athanasius F. M. Marée", "Verônica A. Grieneisen", "Leah Edelstein-Keshet"], "doi"=>["https://dx.doi.org/10.1371/journal.pcbi.1002402.g005"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Feedback_of_cell_shape_dynamics_on_intracellular_dynamics_/343564", "title"=>"Feedback of cell shape dynamics on intracellular dynamics.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2012-03-01 00:59:24"}
  • {"files"=>["https://ndownloader.figshare.com/files/673174"], "description"=>"<p>Effect of phosphoinositide feedback () versus absent feedback () and cell shape dynamics on the response of cells to competing, conflicting, or noisy stimuli. (a) “V” shaped gradient with static cell shape. (See <a href=\"http://www.ploscompbiol.org/article/info:doi/10.1371/journal.pcbi.1002402#pcbi.1002402.s006\" target=\"_blank\">Video S6</a>). (b) “V” shaped gradient with dynamic shape changes. (See <a href=\"http://www.ploscompbiol.org/article/info:doi/10.1371/journal.pcbi.1002402#pcbi.1002402.s007\" target=\"_blank\">Video S7</a>). (c) Noise-induced fronts within motile cells. (See <a href=\"http://www.ploscompbiol.org/article/info:doi/10.1371/journal.pcbi.1002402#pcbi.1002402.s008\" target=\"_blank\">Video S8</a>). PIs allow the cell to rapidly resolve the competition between contradictory signals.</p>", "links"=>[], "tags"=>["Computational biology", "biophysics"], "article_id"=>343670, "categories"=>["Biological Sciences", "Biophysics"], "users"=>["Athanasius F. M. Marée", "Verônica A. Grieneisen", "Leah Edelstein-Keshet"], "doi"=>["https://dx.doi.org/10.1371/journal.pcbi.1002402.g006"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Resolving_conflicts_/343670", "title"=>"Resolving conflicts.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2012-03-01 01:01:10"}
  • {"files"=>["https://ndownloader.figshare.com/files/673302"], "description"=>"<p>Cells initially polarized and moving rightwards encounter a wall (blue vertical band). (<b>a</b>) without feedback from PIs () the cell has difficulty resolving the competition between two nascent “fronts”; (<b>b</b>) with the standard intermediate feedback from PIs () the cell is able to make a decision and move smoothly along the wall; (<b>c</b>) with extreme feedback from PIs () the leading edge becomes confined and encircled by the contracting back, so that the cell becomes stuck. (See also <a href=\"http://www.ploscompbiol.org/article/info:doi/10.1371/journal.pcbi.1002402#pcbi.1002402.s009\" target=\"_blank\">Video S9</a>.)</p>", "links"=>[], "tags"=>["cells", "varying", "pi"], "article_id"=>343795, "categories"=>["Biological Sciences", "Biophysics"], "users"=>["Athanasius F. M. Marée", "Verônica A. Grieneisen", "Leah Edelstein-Keshet"], "doi"=>["https://dx.doi.org/10.1371/journal.pcbi.1002402.g007"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Response_of_cells_with_varying_degree_of_PI_feedback_to_a_wall_/343795", "title"=>"Response of cells with varying degree of PI feedback to a wall.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2012-03-01 01:03:15"}
  • {"files"=>["https://ndownloader.figshare.com/files/673403"], "description"=>"<p>As in <a href=\"http://www.ploscompbiol.org/article/info:doi/10.1371/journal.pcbi.1002402#pcbi-1002402-g007\" target=\"_blank\">Fig. 7</a>, the cell is initially polarized and moving to the right until it encounters an obstacle (static green circular object). (<b>a</b>) Without feedback from PIs () the cell has difficulty resolving the competition between two nascent “fronts” which embrace the object. (<b>b</b>) The cell with feedback from PIs (). The cell is able to move around the object smoothly. (See also <a href=\"http://www.ploscompbiol.org/article/info:doi/10.1371/journal.pcbi.1002402#pcbi.1002402.s010\" target=\"_blank\">Video S10</a>.)</p>", "links"=>[], "tags"=>["pis", "obstacles"], "article_id"=>343896, "categories"=>["Biological Sciences", "Biophysics"], "users"=>["Athanasius F. M. Marée", "Verônica A. Grieneisen", "Leah Edelstein-Keshet"], "doi"=>["https://dx.doi.org/10.1371/journal.pcbi.1002402.g008"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Response_of_a_cell_with_and_without_PIs_to_obstacles_in_its_path_/343896", "title"=>"Response of a cell with and without PIs to obstacles in its path.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2012-03-01 01:04:56"}
  • {"files"=>["https://ndownloader.figshare.com/files/673470"], "description"=>"<p><b>(left)</b> Filament and barbed end densities are described at each grid point within the simulated cell (schematically shown by two representative hexagons), and for all possible orientations , shown at the bottom left. Additionally, close to the membrane we specifically distinguish between barbed ends that have not (yet) reached the membrane () and barbed ends that are effectively pushing against the membrane (), thereby contributing to the forces required for cell extension. <b>(right)</b> The CPM allows for cell shape changes and movement through updates corresponding to small site extensions and retractions. Here, these updates take into account the density of pushing barbed ends. When the cell extends, the pushing barbed ends increase the likelihood of extension (top right – note that all red filaments end up with a pushing barbed end and therefore contribute to the forward motion), while during retraction the barbed ends offer resistence, reducing the likelihood of retraction (bottom right – also note that many black (non force-bearing) filaments are promoted to red (force-bearing) filaments when the retraction is accepted).</p>", "links"=>[], "tags"=>["extensions"], "article_id"=>343967, "categories"=>["Biological Sciences", "Biophysics"], "users"=>["Athanasius F. M. Marée", "Verônica A. Grieneisen", "Leah Edelstein-Keshet"], "doi"=>["https://dx.doi.org/10.1371/journal.pcbi.1002402.g009"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Membrane_extensions_and_retractions_/343967", "title"=>"Membrane extensions and retractions.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2012-03-01 01:06:07"}
  • {"files"=>["https://ndownloader.figshare.com/files/673558"], "description"=>"<p>Note citations of our earlier works (i.e. <a href=\"http://www.ploscompbiol.org/article/info:doi/10.1371/journal.pcbi.1002402#pcbi.1002402-Dawes1\" target=\"_blank\">[25]</a>, <a href=\"http://www.ploscompbiol.org/article/info:doi/10.1371/journal.pcbi.1002402#pcbi.1002402-Mare1\" target=\"_blank\">[36]</a>, <a href=\"http://www.ploscompbiol.org/article/info:doi/10.1371/journal.pcbi.1002402#pcbi.1002402-Jilkine2\" target=\"_blank\">[91]</a>–<a href=\"http://www.ploscompbiol.org/article/info:doi/10.1371/journal.pcbi.1002402#pcbi.1002402-Dawes3\" target=\"_blank\">[93]</a>), where many detailed derivations, explanations, and references can be found.</p>", "links"=>[], "tags"=>["estimates", "actin", "phosphoinositide", "sources"], "article_id"=>344047, "categories"=>["Biological Sciences", "Biophysics"], "users"=>["Athanasius F. M. Marée", "Verônica A. Grieneisen", "Leah Edelstein-Keshet"], "doi"=>["https://dx.doi.org/10.1371/journal.pcbi.1002402.t001"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Parameter_estimates_relevant_to_i_actin_dynamics_ii_Rho_proteins_iii_phosphoinositide_dynamics_and_iv_cell_surface_mechanics_with_sources_from_which_they_were_obtained_/344047", "title"=>"Parameter estimates relevant to (i) actin dynamics; (ii) Rho-proteins; (iii) phosphoinositide dynamics; and (iv) cell surface mechanics, with sources from which they were obtained.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2012-03-01 01:07:27"}

PMC Usage Stats | Further Information

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  • {"unique-ip"=>"6", "full-text"=>"5", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"9"}
  • {"unique-ip"=>"10", "full-text"=>"9", "pdf"=>"4", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"10", "cited-by"=>"0", "year"=>"2016", "month"=>"10"}
  • {"unique-ip"=>"6", "full-text"=>"7", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"2", "supp-data"=>"10", "cited-by"=>"1", "year"=>"2016", "month"=>"11"}
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  • {"unique-ip"=>"5", "full-text"=>"3", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"1", "year"=>"2017", "month"=>"8"}
  • {"unique-ip"=>"2", "full-text"=>"1", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"9"}
  • {"unique-ip"=>"5", "full-text"=>"3", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"10"}
  • {"unique-ip"=>"9", "full-text"=>"8", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"3", "cited-by"=>"1", "year"=>"2017", "month"=>"11"}
  • {"unique-ip"=>"7", "full-text"=>"5", "pdf"=>"5", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"12"}
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  • {"unique-ip"=>"19", "full-text"=>"21", "pdf"=>"3", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"0", "cited-by"=>"1", "year"=>"2018", "month"=>"4"}
  • {"unique-ip"=>"21", "full-text"=>"89", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"1", "year"=>"2018", "month"=>"6"}
  • {"unique-ip"=>"8", "full-text"=>"4", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"2", "cited-by"=>"1", "year"=>"2018", "month"=>"7"}
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  • {"unique-ip"=>"9", "full-text"=>"8", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"1", "year"=>"2018", "month"=>"9"}
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  • {"unique-ip"=>"13", "full-text"=>"13", "pdf"=>"2", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"1", "year"=>"2019", "month"=>"8"}
  • {"unique-ip"=>"17", "full-text"=>"20", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"1", "year"=>"2019", "month"=>"9"}
  • {"unique-ip"=>"14", "full-text"=>"15", "pdf"=>"1", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2019", "month"=>"10"}
  • {"unique-ip"=>"16", "full-text"=>"14", "pdf"=>"6", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"1", "year"=>"2019", "month"=>"12"}
  • {"unique-ip"=>"14", "full-text"=>"12", "pdf"=>"2", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"5", "year"=>"2020", "month"=>"2"}
  • {"unique-ip"=>"15", "full-text"=>"12", "pdf"=>"7", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"1", "year"=>"2020", "month"=>"3"}
  • {"unique-ip"=>"11", "full-text"=>"15", "pdf"=>"3", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"1", "year"=>"2020", "month"=>"4"}
  • {"unique-ip"=>"7", "full-text"=>"10", "pdf"=>"2", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2020", "month"=>"5"}
  • {"unique-ip"=>"6", "full-text"=>"5", "pdf"=>"2", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2020", "month"=>"6"}
  • {"unique-ip"=>"1", "full-text"=>"1", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2020", "month"=>"7"}
  • {"unique-ip"=>"7", "full-text"=>"8", "pdf"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"10", "cited-by"=>"0", "year"=>"2020", "month"=>"8"}

Relative Metric

{"start_date"=>"2012-01-01T00:00:00Z", "end_date"=>"2012-12-31T00:00:00Z", "subject_areas"=>[{"subject_area"=>"/Biology and life sciences/Biochemistry", "average_usage"=>[316, 541, 663, 766, 856, 950, 1041, 1128, 1218, 1302, 1382, 1456, 1526, 1593, 1657, 1729, 1796, 1862, 1930, 1999, 2065, 2132, 2202, 2261, 2319]}, {"subject_area"=>"/Physical sciences/Chemistry", "average_usage"=>[302, 508, 622, 720, 804, 888, 973, 1054, 1141, 1219, 1299, 1370, 1442, 1511, 1574, 1644, 1711, 1782, 1846, 1911, 1971, 2030, 2097, 2155, 2217]}]}
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