Emergence of Large-Scale Cell Morphology and Movement from Local Actin Filament Growth Dynamics
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{"title"=>"Emergence of large-scale cell morphology and movement from local actin filament growth dynamics", "type"=>"journal", "authors"=>[{"first_name"=>"Catherine I.", "last_name"=>"Lacayo", "scopus_author_id"=>"14628948500"}, {"first_name"=>"Zachary", "last_name"=>"Pincus", "scopus_author_id"=>"10339920500"}, {"first_name"=>"Martijn M.", "last_name"=>"VanDuijn", "scopus_author_id"=>"56615788700"}, {"first_name"=>"Cyrus A.", "last_name"=>"Wilson", "scopus_author_id"=>"57199052839"}, {"first_name"=>"Daniel A.", "last_name"=>"Fletcher", "scopus_author_id"=>"7202087947"}, {"first_name"=>"Frank B.", "last_name"=>"Gertler", "scopus_author_id"=>"7003755623"}, {"first_name"=>"Alex", "last_name"=>"Mogilner", "scopus_author_id"=>"7004186241"}, {"first_name"=>"Julie A.", "last_name"=>"Theriot", "scopus_author_id"=>"7005380857"}], "year"=>2007, "source"=>"PLoS Biology", "identifiers"=>{"scopus"=>"2-s2.0-34548475260", "sgr"=>"34548475260", "issn"=>"15457885", "doi"=>"10.1371/journal.pbio.0050233", "pmid"=>"17760506", "isbn"=>"1545-7885 (Electronic)\\n1544-9173 (Linking)", "pui"=>"47416600"}, "id"=>"ec50159f-204b-3bbd-80b0-ae278f936844", "abstract"=>"Variations in cell migration and morphology are consequences of changes in underlying cytoskeletal organization and dynamics. We investigated how these large-scale cellular events emerge as direct consequences of small-scale cytoskeletal molecular activities. Because the properties of the actin cytoskeleton can be modulated by actin-remodeling proteins, we quantitatively examined how one such family of proteins, enabled/vasodilator-stimulated phosphoprotein (Ena/VASP), affects the migration and morphology of epithelial fish keratocytes. Keratocytes generally migrate persistently while exhibiting a characteristic smooth-edged \"canoe\" shape, but may also exhibit less regular morphologies and less persistent movement. When we observed that the smooth-edged canoe keratocyte morphology correlated with enrichment of Ena/VASP at the leading edge, we mislocalized and overexpressed Ena/VASP proteins and found that this led to changes in the morphology and movement persistence of cells within a population. Thus, local changes in actin filament dynamics due to Ena/VASP activity directly caused changes in cell morphology, which is coupled to the motile behavior of keratocytes. We also characterized the range of natural cell-to-cell variation within a population by using measurable morphological and behavioral features--cell shape, leading-edge shape, filamentous actin (F-actin) distribution, cell speed, and directional persistence--that we have found to correlate with each other to describe a spectrum of coordinated phenotypes based on Ena/VASP enrichment at the leading edge. This spectrum stretched from smooth-edged, canoe-shaped keratocytes--which had VASP highly enriched at their leading edges and migrated fast with straight trajectories--to more irregular, rounder cells migrating slower with less directional persistence and low levels of VASP at their leading edges. We developed a mathematical model that accounts for these coordinated cell-shape and behavior phenotypes as large-scale consequences of kinetic contributions of VASP to actin filament growth and protection from capping at the leading edge. This work shows that the local effects of actin-remodeling proteins on cytoskeletal dynamics and organization can manifest as global modifications of the shape and behavior of migrating cells and that mathematical modeling can elucidate these large-scale cell behaviors from knowledge of detailed multiscale protein interactions.", "link"=>"http://www.mendeley.com/research/emergence-largescale-cell-morphology-movement-local-actin-filament-growth-dynamics", "reader_count"=>177, "reader_count_by_academic_status"=>{"Unspecified"=>4, "Professor > Associate Professor"=>19, "Researcher"=>52, "Student > Doctoral Student"=>4, "Student > Ph. D. 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Student"=>57, "Student > Postgraduate"=>7, "Other"=>5, "Student > Master"=>6, "Student > Bachelor"=>6, "Lecturer"=>2, "Lecturer > Senior Lecturer"=>1, "Professor"=>14}, "reader_count_by_subject_area"=>{"Unspecified"=>6, "Agricultural and Biological Sciences"=>94, "Chemical Engineering"=>1, "Chemistry"=>5, "Computer Science"=>3, "Earth and Planetary Sciences"=>1, "Engineering"=>21, "Biochemistry, Genetics and Molecular Biology"=>18, "Materials Science"=>1, "Mathematics"=>5, "Medicine and Dentistry"=>5, "Neuroscience"=>1, "Physics and Astronomy"=>13, "Psychology"=>2, "Immunology and Microbiology"=>1}, "reader_count_by_subdiscipline"=>{"Materials Science"=>{"Materials Science"=>1}, "Medicine and Dentistry"=>{"Medicine and Dentistry"=>5}, "Physics and Astronomy"=>{"Physics and Astronomy"=>13}, "Psychology"=>{"Psychology"=>2}, "Mathematics"=>{"Mathematics"=>5}, "Unspecified"=>{"Unspecified"=>6}, "Chemical Engineering"=>{"Chemical Engineering"=>1}, "Engineering"=>{"Engineering"=>21}, "Chemistry"=>{"Chemistry"=>5}, "Neuroscience"=>{"Neuroscience"=>1}, "Earth and Planetary Sciences"=>{"Earth and Planetary Sciences"=>1}, "Immunology and Microbiology"=>{"Immunology and Microbiology"=>1}, "Agricultural and Biological Sciences"=>{"Agricultural and Biological Sciences"=>94}, "Computer Science"=>{"Computer Science"=>3}, "Biochemistry, Genetics and Molecular Biology"=>{"Biochemistry, Genetics and Molecular Biology"=>18}}, "reader_count_by_country"=>{"Canada"=>1, "United States"=>10, "Japan"=>1, "Brazil"=>1, "United Kingdom"=>5, "Italy"=>1, "France"=>2, "Portugal"=>1, "Germany"=>1}, "group_count"=>7}

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Figshare

  • {"files"=>["https://ndownloader.figshare.com/files/946054"], "description"=>"<div><p>(A, B) Using immunofluorescence, VASP intensity levels were measured along lines (∼5 μm wide) positioned across lamellipodia in the middle of cells roughly perpendicular to the leading edge of keratocytes (arrows). The relative levels of VASP at the leading edge of a population of keratocytes were compared using VASP peak-to-base ratios, which were calculated by dividing the highest mean fluorescence intensity at the leading edge (peak) by the lowest mean intensity found interior to the leading edge (base). For cells with low VASP at the leading edge, peak and base positions were assigned based on mean positions from cells (<i>n</i> = 30) with medium or high VASP peak-to-base ratios and thus clear peak and base positions. (A) An intensity linescan shows that VASP is enriched at the leading edge of a smooth cell with a VASP peak-to-base ratio of 2.56. (B) On the other hand, the rough leading edge of a keratocyte has a VASP peak-to-base ratio of 0.81 and thus VASP is practically absent from the leading edge. Scale bar=10 μm.</p>\n <p>(C) The keratocyte population examined (<i>n =</i> 43) had a wide range of VASP peak-to-base ratio values ranging from 0.5 to 2.8.</p>\n <p>(D) The shapes of keratocytes were compared using PCA, which identified the major modes of shape variation of polygonal cell contours extracted from intensity-thresholded fluorescent images. A shape mode value of zero corresponds to the mean shape and the negative or positive values correspond to standard deviations describing canoe (+SD) or round D shape (−SD) on the <i>y-</i>axis. Representative cell contours for the specified values are shown on the <i>y</i>-axis. For example, the cells in (A) and (B) have shape mode values of 1.32 and −0.77, showing that they can be quantitatively described as canoe or D shaped, respectively. This shape mode significantly correlates with VASP peak-to-base ratios (<i>r<sup>2</sup></i> = 0.28, <i>p</i> = 0.0002, <i>n =</i> 43, solid line).</p>\n <p>(E) To compare leading-edge shapes, we measured and normalized their degree of local curvature. Local leading-edge curvature negatively correlates (<i>r<sup>2</sup></i> = 0.28, <i>p</i> = 0.0003, <i>n =</i> 43, solid line) with VASP peak-to-base ratios.</p>\n <p>(F) A significant negative correlation is also observed between cell shape and local leading-edge curvature (<i>r<sup>2</sup></i> = 0.28, <i>p</i> = 0.0003, <i>n =</i> 43, solid line).</p></div>", "links"=>[], "tags"=>["vasp", "canoe-shaped"], "article_id"=>616417, "categories"=>["Cell Biology", "Biophysics"], "users"=>["Catherine I Lacayo", "Zachary Pincus", "Martijn M VanDuijn", "Cyrus A Wilson", "Daniel A Fletcher", "Frank B Gertler", "Alex Mogilner", "Julie A Theriot"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.0050233.g003", "stats"=>{"downloads"=>1, "page_views"=>9, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Keratocytes_with_High_VASP_at_the_Leading_Edge_Are_Canoe_Shaped_with_Smooth_Leading_Edges_/616417", "title"=>"Keratocytes with High VASP at the Leading Edge Are Canoe-Shaped with Smooth Leading Edges", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2007-08-28 01:46:57"}
  • {"files"=>["https://ndownloader.figshare.com/files/946165"], "description"=>"<p>Outlines of migrating keratocytes overexpressing EGFP-VASP were generated from each frame of time-lapse image sequences separated by 10 s to examine the shape of cells as they migrate. Outlines are colored from blue to red to represent time (0–240 s), superimposed, and plotted on the same scale for visual comparison. Speed can be estimated from the distances traveled by each keratocyte because outlines correspond to the same total time. Fluorescent images correspond to the first (A) and fifth (B) frames of each time-lapse sequence and are scaled to match to the outlines. (A) The leading edge and overall shape of a smooth, coherent keratocyte does not vary extensively as the cell migrates. (B) In the case of a rough and rounder keratocyte, outlines show that the leading edge changes shape rapidly and widely. This keratocyte is unable to maintain persistent coordinated protrusion of its lamellipodium. Specific segments of the leading edge extend forward while adjacent regions lag behind (notice blue and orange outlines). This keratocyte migrates at approximately half the speed of the coherent keratocyte in (A)<i>.</i></p>", "links"=>[], "tags"=>["canoe-shaped", "keratocytes", "migrate", "speeds", "persistent"], "article_id"=>616519, "categories"=>["Cell Biology", "Biophysics"], "users"=>["Catherine I Lacayo", "Zachary Pincus", "Martijn M VanDuijn", "Cyrus A Wilson", "Daniel A Fletcher", "Frank B Gertler", "Alex Mogilner", "Julie A Theriot"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.0050233.g004", "stats"=>{"downloads"=>1, "page_views"=>3, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Smooth_Canoe_Shaped_Keratocytes_Migrate_8220_Coherently_8221_with_Fast_Speeds_and_Persistent_Shapes_/616519", "title"=>"Smooth, Canoe-Shaped Keratocytes Migrate “Coherently,” with Fast Speeds and Persistent Shapes", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2007-08-28 01:48:39"}
  • {"files"=>["https://ndownloader.figshare.com/files/946440"], "description"=>"<div><p>(A) In coherent cells, long actin filaments are protected from capping and undergo significant lateral flow (arrows), smoothing heterogeneities at the leading edge. According to the Graded Radial Extension model, <i>V</i><sub>n</sub>(<i>x</i>)=<i>V</i> cosθ, where <i>V</i> represents cell speed, <i>V</i><sub>n</sub> represents the local protrusion rate, and θ is the orientation of the normal to the leading edge at position <i>x</i>. Quantitative actin dynamics (right) are explained in the <a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.0050233#s4\" target=\"_blank\">Materials and Methods</a> and <a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.0050233#pbio-0050233-sd001\" target=\"_blank\">Text S1</a>.</p>\n <p>(B) In decoherent cells, short filaments that are not protected from capping undergo less-extensive lateral flow (arrows) and may focus into heterogeneities at the leading edge causing the unstable protrusion of microregions.</p>\n <p>(C) Barbed end density and nascent filament branching were chosen so that when VASP activity is low, the F-actin density along the leading edge appears flat (bottom curve). When high VASP activity was entered into our model, the F-actin density along the leading edge emerged as an inverted parabola (top curve), with F-actin density peaked in the middle, as observed experimentally in coherent cells. Position is normalized by the half-length of the leading edge. The prediction that peaked F-actin density is proportional to the level of VASP is qualitatively consistent with the experiment (see <a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.0050233#pbio-0050233-g004\" target=\"_blank\">Figures 4</a>C and <a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.0050233#pbio-0050233-g004\" target=\"_blank\">4</a>D).</p>\n <p>(D) Based on protrusion rate as a function of barbed end distribution along the leading edge, the computed leading edge profile is wide (canoe-shaped) in coherent cells with high VASP at the leading edge (top curve) and short (D shaped) in decoherent cells with low VASP at the leading edge (bottom curve). Position (<i>x</i>) is normalized by the half-length of the leading edge of the short, decoherent cell. The same scale is used for the coherent cell, which is 30% longer, so the corresponding profile extends beyond 1.</p></div>", "links"=>[], "tags"=>["modeling", "explains", "coherent", "decoherent", "keratocyte"], "article_id"=>616788, "categories"=>["Cell Biology", "Biophysics"], "users"=>["Catherine I Lacayo", "Zachary Pincus", "Martijn M VanDuijn", "Cyrus A Wilson", "Daniel A Fletcher", "Frank B Gertler", "Alex Mogilner", "Julie A Theriot"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.0050233.g006", "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Mathematical_Modeling_Explains_the_Coherent_and_Decoherent_Keratocyte_Phenotypes_/616788", "title"=>"Mathematical Modeling Explains the Coherent and Decoherent Keratocyte Phenotypes", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2007-08-28 01:53:08"}
  • {"files"=>["https://ndownloader.figshare.com/files/945776"], "description"=>"<p>A population of primary keratocytes is heterogeneous in morphology. (A) Keratocytes can have a smooth leading edge, showing strong VASP immunofluorescence that appears as a thin line at leading edge (arrowheads). (B) Keratocytes may also have rough leading edges with weak or absent VASP at the leading edge. VASP also appears localized at focal adhesions (arrows), which are more apparent in cells with rough leading edges. Immunofluorescence was performed using polyclonal anti-murine VASP antibodies. Scale bar = 10 μm.</p>", "links"=>[], "tags"=>["vasp", "localization", "observed", "keratocytes", "leading-edge"], "article_id"=>616139, "categories"=>["Cell Biology", "Biophysics"], "users"=>["Catherine I Lacayo", "Zachary Pincus", "Martijn M VanDuijn", "Cyrus A Wilson", "Daniel A Fletcher", "Frank B Gertler", "Alex Mogilner", "Julie A Theriot"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.0050233.g001", "stats"=>{"downloads"=>1, "page_views"=>18, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Strong_VASP_Localization_at_the_Leading_Edge_Is_Observed_in_Keratocytes_with_Smooth_Leading_Edge_Morphology_/616139", "title"=>"Strong VASP Localization at the Leading Edge Is Observed in Keratocytes with Smooth Leading-Edge Morphology", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2007-08-28 01:42:19"}
  • {"files"=>["https://ndownloader.figshare.com/files/946594"], "description"=>"<div><p>(A) Time-lapse images show that overexpressed EGFP-VASP, which is enriched at the leading edge before addition of 1.0 μM cytochalasin D (time: 1:00, left panel), becomes weaker at the leading edge ∼2 min after cytochalasin treatment (time: 2:50, middle panel). Cytochalasin D was added at 70 s (∼1.2 min). Approximately 3 min after cytochalasin treatment (time: 4:20, right panel), EGFP-VASP can barely be seen at the leading edge of this keratocyte that begins to exhibit a D shape rather than the original canoe shape. Time = min:s. Scale bar = 10 μm. Fluorescence intensities measured across the leading edge (arrows) quantitatively confirm the enrichment of EGFP-VASP at the leading edge before cytochalasin treatment and its delocalization a few minutes after addition of the drug (bottom graphs).</p>\n <p>(B) The levels of EGFP-VASP at the leading edge and the width of the keratocyte in (A) were compared as a function of time. Cell width (axis perpendicular to migration) was measured from one side of the cell to the other (dotted line in (A)). EGFP-VASP intensity levels were measured inside a circle (4 μm diameter) placed in the middle of the cell at the leading edge (dotted circles in (A)) in each frame of the time-lapse sequence. These two parameters temporally correlate with each other (<i>r<sup>2</sup> =</i> 0.21, <i>p</i> = 0.0009). The levels of EGFP-VASP at the leading edge (thick gray line) dramatically drop ∼2 min after cytochalasin addition (time ∼3 min, dotted line), followed by a decrease in cell width (thin black line).</p>\n <p>(C) The frequencies of VASP peak-to-base ratios obtained from immunofluorescence images of keratocytes treated with cytochalasin D confirm that VASP becomes displaced from the leading edge. The keratocyte population with high VASP peak-to-base ratios observed in wild-type cells (see <a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.0050233#pbio-0050233-g003\" target=\"_blank\">Figure 3</a>C) becomes absent after cytochalasin treatment, leaving only cells with low and medium values.</p>\n <p>(D) Shape mode analysis reveals that cytochalasin treatment (open diamonds) eliminates the population of keratocytes with canoe shapes (>1 in <i>y-</i>axis) compared to control cells (closed diamonds) causing most cells to resemble the mean shape of the population or rounder D shapes. No significant correlation was found between this shape mode and enrichment of VASP at the leading edge (VASP peak-to-base ratio) in cytochalasin treated cells (<i>r<sup>2</sup></i> = 0.09, <i>n =</i> 27, dashed line) compared to control cells.</p>\n <p>(E) Cytochalasin treatment (open diamonds) eliminates the correlation between local leading-edge curvature and VASP enrichment at the leading edge (<i>r<sup>2</sup></i> = 0.0004, <i>n =</i> 27, dashed line) previously observed in control cells (closed diamonds).</p>\n <p>(F) The negative correlation between cell shape and local leading edge curvature observed in control cells (closed diamonds) is abolished in cells treated with cytochalasin (open diamonds, <i>r<sup>2</sup></i> = 0.0009, <i>n =</i> 27, dashed line).</p>\n <p>(G) After cytochalasin treatment, a typical cell has VASP absent from the leading edge (VASP peak-to-base ratio = 0.83) and an F-actin peak ratio of 0.96 corresponding to a flat F-actin distribution along the leading edge. Cytochalasin treatment eliminated cells with peaked F-actin distributions corresponding to high F-actin peak values from our population. In addition, no significant correlation was found between F-actin peak ratios and enrichment of VASP at the leading edge (VASP peak-to-base ratio) in our population of cytochalasin treated cells (<i>r<sup>2</sup></i> = 0.006, <i>n =</i> 27, dashed line) compared to control cells. Scale bar = 10 μm.</p></div>", "links"=>[], "tags"=>["delocalizes", "vasp", "f-actin"], "article_id"=>616941, "categories"=>["Cell Biology", "Biophysics"], "users"=>["Catherine I Lacayo", "Zachary Pincus", "Martijn M VanDuijn", "Cyrus A Wilson", "Daniel A Fletcher", "Frank B Gertler", "Alex Mogilner", "Julie A Theriot"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.0050233.g007", "stats"=>{"downloads"=>1, "page_views"=>12, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Cytochalasin_Delocalizes_VASP_from_the_Leading_Edge_Changing_the_F_Actin_Network_and_Cell_Shape_/616941", "title"=>"Cytochalasin Delocalizes VASP from the Leading Edge, Changing the F-Actin Network and Cell Shape", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2007-08-28 01:55:41"}
  • {"files"=>["https://ndownloader.figshare.com/files/945941"], "description"=>"<div><p>(A) EGFP-AP4-mito (negative control) binds to mitochondria in the cell body, but does not mislocalize VASP, which, by immunofluorescence, appears as a thin line at the leading edge of cells with a smooth morphology (arrowheads).</p>\n <p>(B) EGFP-FP4-mito mislocalizes VASP at the surface of mitochondria thus preventing its function at the leading edge. Weak VASP localization was only observed at the leading edge in two cells out of more than 50 cells examined. Scale bar = 10 μm.</p>\n <p>(C) Keratocytes were classified as either having a smooth or rough leading edge within populations of migrating keratocytes expressing the aforementioned EGFP-tagged constructs. Cells with a rough leading-edge morphology are more prevalent than those classified as having a smooth morphology when EGFP-FP4-mito is expressed (70%) compared with controls, EGFP (55%) and EGFP-AP4-mito (59%). The incidence of rough keratocytes is significantly lower (43%) when EGFP-VASP is overexpressed (when compared to EGFP-FP4-mito, <i>p</i> < 0.05).</p>\n <p>(D) Keratocytes exhibiting smooth leading edges are significantly faster than those with rough leading-edge morphology (EGFP, <i>p</i> = 0.0002; EGFP-AP4-mito, <i>p =</i> 0.0004; EGFP-FP4-mito, <i>p</i> = 0.0004; EGFP-VASP, <i>p</i> = 0.0065). Mean and SD are plotted.</p>\n <p>(E) To perform qualitative comparisons of keratocyte turning during migration, trajectories of smooth or rough keratocytes were reoriented to start at <i>x,y =</i> 0 in the +<i>y</i> direction in standardized coordinate systems (left and middle panels). Trajectories were truncated (<i>x,y</i> limits = 150 μm) for illustration purposes. The distance between tick marks is 50 μm. For quantitative comparisons, mean angles between velocity vectors separated by specific distances traveled by cells were plotted. Larger mean angles correspond to increased curvature in the trajectories of migrating cells (right panels). Mean angles are significantly smaller for smooth keratocytes expressing negative control constructs compared to rough cells (EGFP, <i>p</i> = 0.0003; EGFP-AP4-mito, <i>p =</i> 0.001), showing that smooth keratocytes maintained straighter paths than rough ones. Ena/VASP protein mislocalization using EGFP-FP4-mito causes smooth cells to move in more curved trajectories with larger mean angles not significantly different from those of rough cells and significantly different from those of smooth cells expressing control constructs (compared to EGFP smooth, <i>p =</i> 0.01; EGFP-AP4-mito smooth, <i>p</i> = 0.005). On the other hand, when keratocytes overexpress EGFP-VASP, rough keratocytes, which have increased trajectory curvature in controls, have straighter paths with smaller angles similar to those from smooth cells expressing controls and EGFP-VASP. Mean and SEM are plotted.</p></div>", "links"=>[], "tags"=>["mislocalization", "decreases", "prevalence", "persistent-moving"], "article_id"=>616302, "categories"=>["Cell Biology", "Biophysics"], "users"=>["Catherine I Lacayo", "Zachary Pincus", "Martijn M VanDuijn", "Cyrus A Wilson", "Daniel A Fletcher", "Frank B Gertler", "Alex Mogilner", "Julie A Theriot"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.0050233.g002", "stats"=>{"downloads"=>1, "page_views"=>10, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Ena_VASP_Mislocalization_Decreases_the_Prevalence_of_Fast_Persistent_Moving_Smooth_Cells_/616302", "title"=>"Ena/VASP Mislocalization Decreases the Prevalence of Fast, Persistent-Moving Smooth Cells", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2007-08-28 01:45:02"}
  • {"files"=>["https://ndownloader.figshare.com/files/946764"], "description"=>"<p>We developed a method to generate nascent lamellipodia during recovery of protrusion after temporarily stalling a section of the lamellipodium. A glass micropipette, acting as a barrier, was lowered into the path of movement of a migrating keratocyte overexpressing EGFP-VASP and forced down until a section flexed parallel to the surface. The pipette was left in place until the cell was in firm contact with the pipette and subsequently removed by translating the pipette in the direction of cell migration. A coherent keratocyte shows EGFP-VASP as a uniform thin line at the leading (time: 0:00, min:s). Fluorescent images are shown in both regular and inverted contrast for clarity. Insets show inversed and zoomed in images of the corresponding boxed areas of the leading edge. When this keratocyte reaches the edge of the barrier, the lamellipodium temporarily stops protruding forward and acquires a very flat shape corresponding to the shape of the barrier (time: 0:48, 0:57). The cell continues migrating in the original direction of motion while EGFP-VASP becomes displaced from the edge of the region in contact with the barrier (see inset). When the micropipette barrier is removed (between 0:57 and 1:00), the leading edge of the lamellipodium immediately resumes protrusion and appears rough with several protruding microregions enriched in EGFP-VASP (time: 1:00). The levels of EGFP-VASP quickly recover along the impacted region and become uniform (time: 1:06). Only 18 s after removal of the barrier, the keratocyte's original shape and EGFP-VASP localization at the leading edge are restored (time: 1:18). Scale bar = 20 μm.</p>", "links"=>[], "tags"=>["lamellipodia", "edges", "accumulating", "vasp", "microprotrusions", "maturing"], "article_id"=>617108, "categories"=>["Cell Biology", "Biophysics"], "users"=>["Catherine I Lacayo", "Zachary Pincus", "Martijn M VanDuijn", "Cyrus A Wilson", "Daniel A Fletcher", "Frank B Gertler", "Alex Mogilner", "Julie A Theriot"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.0050233.g008", "stats"=>{"downloads"=>0, "page_views"=>5, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Nascent_Lamellipodia_Emerge_with_Rough_Edges_Accumulating_VASP_in_Microprotrusions_Eventually_Maturing_into_a_Smooth_Morphology_/617108", "title"=>"Nascent Lamellipodia Emerge with Rough Edges Accumulating VASP in Microprotrusions Eventually Maturing into a Smooth Morphology", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2007-08-28 01:58:28"}
  • {"files"=>["https://ndownloader.figshare.com/files/946337"], "description"=>"<div><p>(A) The distributions of VASP (gray line in the graph) and F-actin (black line in the graph) were measured along the length of the leading edge of cells (position indicated by the arrow). The cell shown has VASP enriched at its smooth leading edge (VASP peak-to-base ratio = 1.84). The distribution of F-actin along the leading edge of this smooth cell is peaked in the middle and strongly correlates with that of VASP (<i>r<sup>2</sup></i> = 0.82, <i>p</i> < 0.0001). Scale bar = 20 μm.</p>\n <p>(B) A keratocyte with rough leading edge and very weak VASP at the leading edge (VASP peak-to-base ratio = 0.76) has a very flat distribution of VASP (gray line in the graph) and F-actin (solid black line in the graph) measured along the leading edge (arrow). The distributions of VASP and F-actin along the leading edge of this cell strongly correlate with each other (<i>r<sup>2</sup></i> = 0.81, <i>p</i> < 0.0001). Additionally, the distributions of VASP and F-actin of the entire population of keratocytes strongly correlate (<<i>r></i> = 0.71, SD = 0.23, <i>p</i> < 0.0001, <i>n</i> = 43).</p>\n <p>(C) The F-actin density along the leading edge of a representative coherent cell with high VASP peak-to-base ratio—2.79 (gray line)—is increased compared to that of a decoherent cell with low VASP peak-to-base ratio—1.13 (black line). Mean intensity values are not normalized and were obtained from keratocytes imaged from the same coverslip. F-actin distributions between different cells were compared by calculating a ratio (F-actin peak ratio) of the mean F-actin intensity values from the middle half of the leading edge (0.25 to 0.75 position along the edge, indicated by the thick regions of each line in the graph), which generally correspond to the highest intensity values in peaked F-actin distributions, to the mean of the F-actin values from the rest of the leading edge (positions 0 to 0.25 and 0.75 to 1.00 along the edge, indicated by the thin regions of each line). Cells with peaked F-actin distributions had larger F-actin peak ratios than did cells with flat distributions. The F-actin peak ratio of the coherent cell (gray line) is 1.45, whereas that of the decoherent cell (black line) is 0.94. Also compare the cells in (A) and<b> (</b>B), which have F-actin peak ratios of 1.56 and 0.94, respectively.</p>\n <p>(D) VASP peak-to-base ratios significantly correlate with F-actin peak ratios (<i>r<sup>2</sup></i> = 0.31, <i>p</i> = 0.0001, <i>n =</i> 43, solid line).</p>\n <p>(E, F) For comparisons of F-actin and Arp3, mean intensity values were measured along lines (∼0.5 μm wide) positioned along the leading edge of keratocytes, immediately interior to cell edge. Anti-Arp3 mean fluorescence intensities measured along the leading edge of keratocytes are consistent with those of F-actin. A representative smooth cell (E) exhibits peaked Arp3 and F-actin distributions along the leading edge while a rough cell (F) has flat distributions. Scale bar = 10 μm.</p></div>", "links"=>[], "tags"=>["enrichment", "correlates", "peaked", "f-actin"], "article_id"=>616692, "categories"=>["Cell Biology", "Biophysics"], "users"=>["Catherine I Lacayo", "Zachary Pincus", "Martijn M VanDuijn", "Cyrus A Wilson", "Daniel A Fletcher", "Frank B Gertler", "Alex Mogilner", "Julie A Theriot"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.0050233.g005", "stats"=>{"downloads"=>1, "page_views"=>4, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_VASP_Enrichment_at_the_Leading_Edge_Correlates_with_Peaked_F_Actin_Distributions_/616692", "title"=>"VASP Enrichment at the Leading Edge Correlates with Peaked F-Actin Distributions", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2007-08-28 01:51:32"}
  • {"files"=>["https://ndownloader.figshare.com/files/466161", "https://ndownloader.figshare.com/files/466220", "https://ndownloader.figshare.com/files/466267", "https://ndownloader.figshare.com/files/466327", "https://ndownloader.figshare.com/files/466503", "https://ndownloader.figshare.com/files/466544"], "description"=>"<div><p>Variations in cell migration and morphology are consequences of changes in underlying cytoskeletal organization and dynamics. We investigated how these large-scale cellular events emerge as direct consequences of small-scale cytoskeletal molecular activities. Because the properties of the actin cytoskeleton can be modulated by actin-remodeling proteins, we quantitatively examined how one such family of proteins, enabled/vasodilator-stimulated phosphoprotein (Ena/VASP), affects the migration and morphology of epithelial fish keratocytes. Keratocytes generally migrate persistently while exhibiting a characteristic smooth-edged “canoe” shape, but may also exhibit less regular morphologies and less persistent movement. When we observed that the smooth-edged canoe keratocyte morphology correlated with enrichment of Ena/VASP at the leading edge, we mislocalized and overexpressed Ena/VASP proteins and found that this led to changes in the morphology and movement persistence of cells within a population. Thus, local changes in actin filament dynamics due to Ena/VASP activity directly caused changes in cell morphology, which is coupled to the motile behavior of keratocytes. We also characterized the range of natural cell-to-cell variation within a population by using measurable morphological and behavioral features—cell shape, leading-edge shape, filamentous actin (F-actin) distribution, cell speed, and directional persistence—that we have found to correlate with each other to describe a spectrum of coordinated phenotypes based on Ena/VASP enrichment at the leading edge. This spectrum stretched from smooth-edged, canoe-shaped keratocytes—which had VASP highly enriched at their leading edges and migrated fast with straight trajectories—to more irregular, rounder cells migrating slower with less directional persistence and low levels of VASP at their leading edges. We developed a mathematical model that accounts for these coordinated cell-shape and behavior phenotypes as large-scale consequences of kinetic contributions of VASP to actin filament growth and protection from capping at the leading edge. This work shows that the local effects of actin-remodeling proteins on cytoskeletal dynamics and organization can manifest as global modifications of the shape and behavior of migrating cells and that mathematical modeling can elucidate these large-scale cell behaviors from knowledge of detailed multiscale protein interactions.</p> </div>", "links"=>[], "tags"=>["emergence", "large-scale", "morphology", "actin", "filament"], "article_id"=>151892, "categories"=>["Cell Biology", "Biophysics"], "users"=>["Catherine I Lacayo", "Zachary Pincus", "Martijn M VanDuijn", "Cyrus A Wilson", "Daniel A Fletcher", "Frank B Gertler", "Alex Mogilner", "Julie A Theriot"], "doi"=>["https://dx.doi.org/10.1371/journal.pbio.0050233.sg001", "https://dx.doi.org/10.1371/journal.pbio.0050233.sg002", "https://dx.doi.org/10.1371/journal.pbio.0050233.sg003", "https://dx.doi.org/10.1371/journal.pbio.0050233.sd001", "https://dx.doi.org/10.1371/journal.pbio.0050233.sv001", "https://dx.doi.org/10.1371/journal.pbio.0050233.sv002"], "stats"=>{"downloads"=>6, "page_views"=>9, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/Emergence_of_Large_Scale_Cell_Morphology_and_Movement_from_Local_Actin_Filament_Growth_Dynamics/151892", "title"=>"Emergence of Large-Scale Cell Morphology and Movement from Local Actin Filament Growth Dynamics", "pos_in_sequence"=>0, "defined_type"=>4, "published_date"=>"2007-08-28 00:31:32"}

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  • {"unique-ip"=>"10", "full-text"=>"8", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"5", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2015", "month"=>"9"}
  • {"unique-ip"=>"6", "full-text"=>"10", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"1", "cited-by"=>"0", "year"=>"2015", "month"=>"10"}
  • {"unique-ip"=>"7", "full-text"=>"4", "pdf"=>"3", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"3", "supp-data"=>"0", "cited-by"=>"5", "year"=>"2015", "month"=>"11"}
  • {"unique-ip"=>"13", "full-text"=>"9", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"3", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2015", "month"=>"12"}
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  • {"unique-ip"=>"8", "full-text"=>"8", "pdf"=>"5", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"3", "cited-by"=>"0", "year"=>"2016", "month"=>"10"}
  • {"unique-ip"=>"7", "full-text"=>"7", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"3", "cited-by"=>"0", "year"=>"2016", "month"=>"11"}
  • {"unique-ip"=>"8", "full-text"=>"7", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"3", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2016", "month"=>"12"}
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  • {"unique-ip"=>"8", "full-text"=>"9", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"2", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"2"}
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  • {"unique-ip"=>"7", "full-text"=>"6", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"2", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"9"}
  • {"unique-ip"=>"2", "full-text"=>"3", "pdf"=>"1", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"10"}
  • {"unique-ip"=>"7", "full-text"=>"10", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"9", "supp-data"=>"1", "cited-by"=>"0", "year"=>"2017", "month"=>"11"}
  • {"unique-ip"=>"3", "full-text"=>"6", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2017", "month"=>"12"}
  • {"unique-ip"=>"6", "full-text"=>"9", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"2", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"1"}
  • {"unique-ip"=>"1", "full-text"=>"1", "pdf"=>"0", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"0", "supp-data"=>"0", "cited-by"=>"0", "year"=>"2018", "month"=>"2"}
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Relative Metric

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