316:
According to this model, microdomain signaling dynamics organizes cytoskeleton and its interaction with substratum. As microdomains trigger and maintain active polymerization of actin filaments, their propagation and zigzagging motion on the membrane generate a highly interlinked network of curved or linear filaments oriented at a wide spectrum of angles to the cell boundary. It is also proposed that microdomain interaction marks the formation of new focal adhesion sites at the cell periphery. Myosin interaction with the actin network then generate membrane retraction/ruffling, retrograde flow, and contractile forces for forward motion. Finally, continuous application of stress on the old focal adhesion sites could result in the calcium-induced calpain activation, and consequently the detachment of focal adhesions which completes the cycle.
218:
are dynamic, they are able to remodel to allow retraction. When dynamics are suppressed, microtubules cannot remodel and, therefore, oppose the contractile forces. The morphology of cells with suppressed microtubule dynamics indicate that cells can extend the front edge (polarized in the direction of movement), but have difficulty retracting their trailing edge. On the other hand, high drug concentrations, or microtubule mutations that depolymerize the microtubules, can restore cell migration but there is a loss of directionality. It can be concluded that microtubules act both to restrain cell movement and to establish directionality.
328:βa front and a back. Without it, they would move in all directions at once, i.e. spread. How this polarity is formulated at a molecular level inside a cell is unknown. In a cell that is meandering in a random way, the front can easily give way to become passive as some other region, or regions, of the cell form(s) a new front. In chemotaxing cells, the stability of the front appears enhanced as the cell advances toward a higher concentration of the stimulating chemical. From biophysical perspective, polarity was explained in terms of a gradient in inner membrane
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of paramount importance. The mathematical models developed in these works determine some physical features and material properties of the cells locally through analysis of live cell image sequences and uses this information to make further inferences about the molecular structures, dynamics, and processes within the cells, such as the actin network, microdomains, chemotaxis, adhesion, and retrograde flow.
171:
230:. This suggests that extension of the leading edge occurs primarily by addition of membrane at the front of the cell. If so, the actin filaments that form there might stabilize the added membrane so that a structured extension, or lamella, is formed β rather than a bubble-like structure (or bleb) at its front. For a cell to move, it is necessary to bring a fresh supply of "feet" (proteins called
398:
294:, that cells migrating in an amoeboid fashion without adhesions exhibit plasma membrane flow towards the cell rear that may propel cells by exerting tangential forces on the surrounding fluid. Polarized trafficking of membrane-containing vesicles from the rear to the front of the cell helps maintain cell size. Rearward membrane flow was also observed in
182:(A) Dynamic microtubules are necessary for tail retraction and are distributed at the rear end in a migrating cell. Green, highly dynamic microtubules; yellow, moderately dynamic microtubules and red, stable microtubules. (B) Stable microtubules act as struts and prevent tail retraction and thereby inhibit cell migration.
372:
Although microtubules have been known to influence cell migration for many years, the mechanism by which they do so has remained controversial. On a planar surface, microtubules are not needed for the movement, but they are required to provide directionality to cell movement and efficient protrusion
315:
Based on some mathematical models, recent studies hypothesize a novel biological model for collective biomechanical and molecular mechanism of cell motion. It is proposed that microdomains weave the texture of cytoskeleton and their interactions mark the location for formation of new adhesion sites.
385:
in cell motility has been established. This approach is based on the idea that behavioral or shape changes of a cell bear information about the underlying mechanisms that generate these changes. Reading cell motion, namely, understanding the underlying biophysical and mechanochemical processes, is
285:
fluid dynamics, which apply at the cellular scale, rearward surface flow could provide a mechanism for microscopic objects to swim forward. After some decades, experimental support for this model of cell movement was provided when it was discovered (in 2010) that amoeboid cells and neutrophils are
217:
Other cytoskeletal components (like microtubules) have important functions in cell migration. It has been found that microtubules act as "struts" that counteract the contractile forces that are needed for trailing edge retraction during cell movement. When microtubules in the trailing edge of cell
208:
polymerisation at the cell's front edge. This observation has led to the hypothesis that formation of actin filaments "push" the leading edge forward and is the main motile force for advancing the cell's front edge. In addition, cytoskeletal elements are able to interact extensively and intimately
120:
videos are recorded of the migrating cells to speed up the movement. Such videos (Figure 1) reveal that the leading cell front is very active, with a characteristic behavior of successive contractions and expansions. It is generally accepted that the leading front is the main motor that pulls the
364:
are transported along these filaments to the cell's front. In chemotaxing cells, the increased persistence of migration toward the target may result from an increased stability of the arrangement of the filamentous structures inside the cell and determine its polarity. In turn, these filamentous
248:
which affect membrane recycling block cell migration at the restrictive (higher) temperature; they provide additional support for the importance of the endocytic cycle in cell migration. Furthermore, these amoebae move quite quickly β about one cell length in ~5 mins. If they are regarded as
298:
cells. These observations provide strong support for models of cell movement which depend on a rearward cell surface membrane flow (Model B, above). Interestingly, the migration of supracellular clusters has also been found to be supported by a similar mechanism of rearward surface flow.
359:
are important for establishing and maintaining a cell's polarity. Drugs that destroy actin filaments have multiple and complex effects, reflecting the wide role that these filaments play in many cell processes. It may be that, as part of the locomotory process, membrane
234:, which attach a cell to the surface on which it is crawling) to the front. It is likely that these feet are endocytosed toward the rear of the cell and brought to the cell's front by exocytosis, to be reused to form new attachments to the substrate.
365:
structures may be arranged inside the cell according to how molecules like PIP3 and PTEN are arranged on the inner cell membrane. And where these are located appears in turn to be determined by the chemoattractant signals as these impinge on specific
191:
There are two main theories for how the cell advances its front edge: the cytoskeletal model and membrane flow model. It is possible that both underlying processes contribute to cell extension.
92:
cell migration typically is far more complex and can consist of combinations of different migration mechanisms. It generally involves drastic changes in cell shape which are driven by the
249:
cylindrical (which is roughly true whilst chemotaxing), this would require them to recycle the equivalent of one cell surface area each 5 mins, which is approximately what is measured.
80:), cells need to continuously produce forces in order to move. Cells achieve active movement by very different mechanisms. Many less complex prokaryotic organisms (and sperm cells) use
2221:
Lin Y, Pal DS, Banerjee P, Banerjee T, Qin G, Deng Y, et al. (July 2024). "Ras suppression potentiates rear actomyosin contractility-driven cell polarization and migration".
73:. An understanding of the mechanism by which cells migrate may lead to the development of novel therapeutic strategies for controlling, for example, invasive tumour cells.
1319:"In migrating fibroblasts, recycling receptors are concentrated in narrow tubules in the pericentriolar area, and then routed to the plasma membrane of the leading lamella"
2981:
332:
between front regions and rear edges of the cell. This polarity is reflected at a molecular level by a restriction of certain molecules to particular regions of the inner
899:
Abercrombie M, Heaysman JE, Pegrum SM (October 1970). "The locomotion of fibroblasts in culture. 3. Movements of particles on the dorsal surface of the leading lamella".
49:
all require the orchestrated movement of cells in particular directions to specific locations. Cells often migrate in response to specific external signals, including
226:
The leading edge at the front of a migrating cell is also the site at which membrane from internal membrane pools is returned to the cell surface at the end of the
100:
motility. A paradigmatic example of crawling motion is the case of fish epidermal keratocytes, which have been extensively used in research and teaching.
337:
303:
2404:
Coskun H, Coskun H (March 2011). "Cell physician: reading cell motion: a mathematical diagnostic technique through analysis of single cell motion".
1974:
Coskun H, Coskun H (March 2011). "Cell physician: reading cell motion: a mathematical diagnostic technique through analysis of single cell motion".
436:
373:
of the leading edge. When present, microtubules retard cell movement when their dynamics are suppressed by drug treatment or by tubulin mutations.
2670:
2850:
2361:
Coskun H, Li Y, Mackey MA (January 2007). "Ameboid cell motility: a model and inverse problem, with an application to live cell imaging data".
2589:
736:"Cell migration analysis: A low-cost laboratory experiment for cell and developmental biology courses using keratocytes from fish scales"
2599:
2486:
934:
Willard SS, Devreotes PN (September 2006). "Signaling pathways mediating chemotaxis in the social amoeba, Dictyostelium discoideum".
2792:
256:
Rearward membrane flow (red arrows) and vesicle trafficking from back to front (blue arrows) drive adhesion-independent migration.
2689:
160:
869:
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2511:
245:
3002:
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2452:
677:"A novel 2.5D culture platform to investigate the role of stiffness gradients on adhesion-independent cell migration"
148:
The latter feature is most easily observed when aggregates of a surface molecule are cross-linked with a fluorescent
2516:
1464:
Thompson CR, Bretscher MS (September 2002). "Cell polarity and locomotion, as well as endocytosis, depend on NSF".
1417:"Circulating integrins: alpha 5 beta 1, alpha 6 beta 4 and Mac-1, but not alpha 3 beta 1, alpha 4 beta 1 or LFA-1"
265:
Adhesive crawling is not the only migration mode exhibited by eukaryotic cells. Importantly, several cell types β
167:
genome that simplifies the process of connecting a particular gene product with its effect on cellular behaviour.
2694:
2665:
2565:
1153:"Inhibition of cell migration and cell division correlates with distinct effects of microtubule inhibiting drugs"
565:"An excitable Ras/PI3K/ERK signaling network controls migration and oncogenic transformation in epithelial cells"
421:
1116:
Doherty GJ, McMahon HT (2008). "Mediation, modulation, and consequences of membrane-cytoskeleton interactions".
2845:
238:
156:
2252:
Li R, Gundersen GG (November 2008). "Beyond polymer polarity: how the cytoskeleton builds a polarized cell".
1501:"Using single loxP sites to enhance homologous recombination: ts mutants in Sec1 of Dictyostelium discoideum"
826:"Live-cell imaging of migrating cells expressing fluorescently-tagged proteins in a three-dimensional matrix"
516:"Eukaryotic Chemotaxis: A Network of Signaling Pathways Controls Motility, Directional Sensing, and Polarity"
2926:
2711:
366:
58:
290:
towards a chemo-attractant source whilst suspended in an isodense medium. It was subsequently shown, using
3007:
2946:
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2479:
1260:"Distribution of receptors for transferrin and low density lipoprotein on the surface of giant HeLa cells"
62:
34:
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1868:
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1719:
1512:
1271:
971:"Exchange of actin subunits at the leading edge of living fibroblasts: possible role of treadmilling"
688:
625:
278:
2297:"2D protrusion but not motility predicts growth factor-induced cancer cell migration in 3D collagen"
2976:
2951:
2941:
2531:
2117:
Ridley AJ, Schwartz MA, Burridge K, Firtel RA, Ginsberg MH, Borisy G, et al. (December 2003).
660:
129:
The processes underlying mammalian cell migration are believed to be consistent with those of (non-
97:
252:
2956:
2817:
2812:
2432:
2277:
2154:
2056:
1999:
1397:
1098:
1049:
1914:"Supracellular contraction at the rear of neural crest cell groups drives collective chemotaxis"
306:
Schematic representation of the collective biomechanical and molecular mechanism of cell motion
178:
277:β have been found to be capable of adhesion-independent migration. Historically, the physicist
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2840:
2827:
2787:
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2628:
2472:
2455:
The Cell
Migration Gateway is a comprehensive and regularly updated resource on cell migration
2421:
2386:
2326:
2295:
Meyer AS, Hughes-Alford SK, Kay JE, Castillo A, Wells A, Gertler FB, et al. (June 2012).
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1991:
1951:
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O'Neill PR, Castillo-Badillo JA, Meshik X, Kalyanaraman V, Melgarejo K, Gautam N (July 2018).
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2174:"Actuation of single downstream nodes in growth factor network steers immune cell migration"
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1983:
1941:
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Regulated self-propelled movement of cells from one site to another guided by molecular cues
2779:
2618:
2503:
2495:
2172:
Pal DS, Banerjee T, Lin Y, de
Trogoff F, Borleis J, Iglesias PA, et al. (July 2023).
2019:"Spatiotemporal dynamics of membrane surface charge regulates cell polarity and migration"
612:
Huber F, SchnauΓ J, RΓΆnicke S, Rauch P, MΓΌller K, FΓΌtterer C, et al. (January 2013).
531:
431:
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227:
77:
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2134:
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2018:
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96:. Two very distinct migration scenarios are crawling motion (most commonly studied) and
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2677:
2570:
2321:
2296:
1946:
1913:
1889:
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1791:
1756:
1687:
1662:
1584:
1560:"The exocytic gene secA is required for Dictyostelium cell motility and osmoregulation"
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1036:
1019:
467:"Single-Cell Migration in Complex Microenvironments: Mechanics and Signaling Dynamics"
174:
Two different models for how cells move. A) Cytoskeletal model. B) Membrane Flow Model
2996:
2835:
2577:
2548:
2526:
2060:
1633:
1608:
912:
563:
Zhan H, Bhattacharya S, Cai H, Iglesias PA, Huang CH, Devreotes PN (September 2020).
411:
333:
325:
159:
is useful to researchers because they consistently exhibit chemotaxis in response to
46:
42:
38:
2281:
2158:
2003:
1401:
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170:
130:
109:
93:
54:
1855:
Tanaka M, Kikuchi T, Uno H, Okita K, Kitanishi-Yumura T, Yumura S (October 2017).
1053:
2433:"Mathematicians use cell 'profiling' to detect abnormalities -- including cancer"
2189:
2095:
1832:
1815:
1678:
1525:
701:
637:
580:
2936:
2885:
2726:
2582:
2017:
Banerjee T, Biswas D, Pal DS, Miao Y, Iglesias PA, Devreotes PN (October 2022).
441:
356:
349:
2382:
2230:
2034:
1880:
1761:
Proceedings of the
National Academy of Sciences of the United States of America
1264:
Proceedings of the
National Academy of Sciences of the United States of America
947:
877:
397:
2743:
2721:
2417:
2347:
Mathematical Models for
Ameboid Cell Motility and Model Based Inverse Problems
2118:
1987:
416:
393:
287:
274:
270:
113:
89:
70:
50:
17:
792:
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2142:
1937:
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426:
231:
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2273:
2238:
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2150:
2103:
2052:
1995:
1955:
1898:
1841:
1800:
1696:
1663:"Membrane Flow Drives an Adhesion-Independent Amoeboid Cell Migration Mode"
1642:
1624:
1593:
1544:
1485:
1334:
1317:
Hopkins CR, Gibson A, Shipman M, Strickland DK, Trowbridge IS (June 1994).
1244:
1188:
1137:
1094:
955:
855:
810:
761:
720:
655:
598:
549:
500:
2312:
1450:
1393:
1352:
1303:
1284:
1045:
1004:
986:
920:
614:"Emergent complexity of the cytoskeleton: from single filaments to tissue"
144:
laminar removal of dorsally-accumulated debris toward trailing edge (back)
2900:
2862:
2594:
2536:
2461:
A tour of images and videos by the J. V. Small lab in
Salzburg and Vienna
1368:"Getting membrane flow and the cytoskeleton to cooperate in moving cells"
1069:"Cellular motility driven by assembly and disassembly of actin filaments"
149:
81:
163:; they move more quickly than cultured mammalian cells; and they have a
152:
or when small beads become artificially bound to the front of the cell.
2521:
1740:
1575:
164:
752:
735:
482:
2555:
2074:
Parent CA, Devreotes PN (April 1999). "A cell's sense of direction".
155:
Other eukaryotic cells are observed to migrate similarly. The amoeba
2265:
1731:
2797:
341:
301:
251:
205:
177:
169:
85:
66:
57:. Errors during this process have serious consequences, including
1207:
Ganguly A, Yang H, Sharma R, Patel KD, Cabral F (December 2012).
837:
2543:
2458:
345:
2468:
1209:"The role of microtubules and their dynamics in cell migration"
311:
Collective biomechanical and molecular mechanism of cell motion
2345:
2464:
1857:"Turnover and flow of the cell membrane for cell migration"
33:
is a central process in the development and maintenance of
734:
Prieto D, Aparicio G, Sotelo-Silveira JR (November 2017).
1558:
Zanchi R, Howard G, Bretscher MS, Kay RR (October 2010).
112:
attached to a surface or in 3D is commonly studied using
2350:(Ph.D. thesis). University of Iowa – via ProQuest.
2119:"Cell migration: integrating signals from front to back"
1912:
Shellard A, SzabΓ³ A, Trepat X, Mayor R (October 2018).
670:
668:
1816:""Rho"ing a Cellular Boat with Rearward Membrane Flow"
1609:"Circulation of the plasma membrane in Dictyostelium"
2909:
2871:
2826:
2778:
2646:
2502:
465:Mak M, Spill F, Kamm RD, Zaman MH (February 2016).
2982:Task allocation and partitioning of social insects
1710:Purcell EM (1977). "Life at Low Reynolds Number".
116:. As cell movement is very slow, a few ΞΌm/minute,
281:theorized (in 1977) that under conditions of low
1757:"Dictyostelium amoebae and neutrophils can swim"
1607:Aguado-Velasco C, Bretscher MS (December 1999).
377:Inverse problems in the context of cell motility
141:cytoplasmic displacement at leading edge (front)
1020:"Actin-based cell motility and cell locomotion"
1656:
1654:
1652:
204:Experimentation has shown that there is rapid
2480:
1969:
1967:
1965:
1202:
1200:
1198:
8:
1151:Yang H, Ganguly A, Cabral F (October 2010).
740:Biochemistry and Molecular Biology Education
355:It is believed that filamentous actins and
344:are found at the front of the cell, whereas
876:. Cell Migration Consortium. Archived from
76:Due to the highly viscous environment (low
2487:
2473:
2465:
1499:Bretscher MS, Clotworthy M (August 2007).
2320:
2197:
2042:
1945:
1888:
1831:
1790:
1780:
1739:
1686:
1632:
1583:
1534:
1524:
1440:
1383:
1342:
1293:
1283:
1234:
1224:
1178:
1168:
1084:
1035:
1018:Mitchison TJ, Cramer LP (February 1996).
994:
845:
800:
751:
710:
700:
645:
588:
539:
514:Swaney K, Huang CH, Devreotes PN (2010).
490:
1130:10.1146/annurev.biophys.37.032807.125912
675:Pebworth MP, Cismas SA, Asuri P (2014).
437:Mouse models of breast cancer metastasis
1067:Pollard TD, Borisy GG (February 2003).
457:
261:Mechanistic basis of amoeboid migration
2254:Nature Reviews. Molecular Cell Biology
2590:Patterns of self-organization in ants
532:10.1146/annurev.biophys.093008.131228
7:
1755:Barry NP, Bretscher MS (June 2010).
775:Dormann D, Weijer CJ (August 2006).
471:Journal of Biomechanical Engineering
2459:The Cytoskeleton and Cell Migration
1213:The Journal of Biological Chemistry
1157:The Journal of Biological Chemistry
137:. Observations in common include:
2600:symmetry breaking of escaping ants
1433:10.1002/j.1460-2075.1992.tb05068.x
824:Shih W, Yamada S (December 2011).
25:
1814:Bell GR, Collins SR (July 2018).
830:Journal of Visualized Experiments
2637:
2406:Bulletin of Mathematical Biology
1976:Bulletin of Mathematical Biology
936:European Journal of Cell Biology
396:
187:Molecular processes of migration
209:with a cell's plasma membrane.
2363:Journal of Theoretical Biology
1415:Bretscher MS (February 1992).
1366:Bretscher MS (November 1996).
273:, metastatic cancer cells and
1:
1613:Molecular Biology of the Cell
1385:10.1016/S0092-8674(00)81380-X
1258:Bretscher MS (January 1983).
1086:10.1016/S0092-8674(03)00120-X
1037:10.1016/S0092-8674(00)81281-7
369:on the cell's outer surface.
246:temperature sensitive mutants
2561:Mixed-species foraging flock
2512:Agent-based model in biology
2190:10.1016/j.devcel.2023.04.019
2096:10.1126/science.284.5415.765
1833:10.1016/j.devcel.2018.06.008
1679:10.1016/j.devcel.2018.05.029
1526:10.1371/journal.pone.0000724
913:10.1016/0014-4827(70)90570-7
702:10.1371/journal.pone.0110453
638:10.1080/00018732.2013.771509
581:10.1016/j.devcel.2020.08.001
340:and activated Ras, Rac, and
2808:Particle swarm optimization
2301:The Journal of Cell Biology
1712:American Journal of Physics
1323:The Journal of Cell Biology
1118:Annual Review of Biophysics
975:The Journal of Cell Biology
777:"Imaging of cell migration"
520:Annual Review of Biophysics
381:An area of research called
352:are found toward the rear.
320:Polarity in migrating cells
3024:
2517:Collective animal behavior
2383:10.1016/j.jtbi.2006.07.025
2231:10.1038/s41556-024-01453-4
2035:10.1038/s41556-022-00997-7
1881:10.1038/s41598-017-13438-5
948:10.1016/j.ejcb.2006.06.003
901:Experimental Cell Research
37:. Tissue formation during
2635:
2418:10.1007/s11538-010-9580-x
1988:10.1007/s11538-010-9580-x
870:"What is Cell Migration?"
422:Collective cell migration
336:. Thus, the phospholipid
2846:Self-propelled particles
793:10.1038/sj.emboj.7601227
296:Dictyostelium discoideum
157:Dictyostelium discoideum
2927:Collective intelligence
2793:Ant colony optimization
2143:10.1126/science.1092053
1938:10.1126/science.aau3301
1782:10.1073/pnas.1006327107
1564:Journal of Cell Science
1478:10.1242/dev.129.18.4185
1226:10.1074/jbc.M112.423905
1170:10.1074/jbc.M110.160820
969:Wang YL (August 1985).
324:Migrating cells have a
222:Membrane flow model (B)
59:intellectual disability
35:multicellular organisms
2947:Microbial intelligence
2607:Shoaling and schooling
2453:Cell Migration Gateway
1625:10.1091/mbc.10.12.4419
1335:10.1083/jcb.125.6.1265
874:Cell Migration Gateway
307:
257:
195:Cytoskeletal model (A)
183:
175:
104:Cell migration studies
88:to propel themselves.
2313:10.1083/jcb.201201003
1285:10.1073/pnas.80.2.454
987:10.1083/jcb.101.2.597
305:
255:
181:
173:
118:time-lapse microscopy
39:embryonic development
2967:Spatial organization
2932:Decentralised system
2770:Sea turtle migration
2624:Swarming (honey bee)
2184:(13): 1170β1188.e7.
1570:(Pt 19): 3226β3234.
244:, three conditional
2942:Group size measures
2504:Biological swarming
2375:2007JThBi.244..169C
2223:Nature Cell Biology
2135:2003Sci...302.1704R
2129:(5651): 1704β1709.
2088:1999Sci...284..765P
2023:Nature Cell Biology
1930:2018Sci...362..339S
1873:2017NatSR...712970T
1773:2010PNAS..10711376B
1767:(25): 11376β11380.
1724:1977AmJPh..45....3P
1517:2007PLoSO...2..724B
1276:1983PNAS...80..454B
1219:(52): 43359β43369.
1163:(42): 32242β32250.
693:2014PLoSO...9k0453P
630:2013AdPhy..62....1H
618:Advances in Physics
3003:Cellular processes
2957:Predator satiation
2818:Swarm (simulation)
2813:Swarm intelligence
2788:Agent-based models
2619:Swarming behaviour
2430:; Lay summary in:
2178:Developmental Cell
1861:Scientific Reports
1820:Developmental Cell
1667:Developmental Cell
1576:10.1242/jcs.072876
880:on 22 October 2014
569:Developmental Cell
308:
258:
184:
176:
55:mechanical signals
2990:
2989:
2977:Military swarming
2922:Animal navigation
2841:Collective motion
2828:Collective motion
2695:reverse migration
2629:Swarming motility
2344:Coskun H (2006).
2082:(5415): 765β770.
2029:(10): 1499β1515.
1924:(6412): 339β343.
1619:(12): 4419β4427.
1472:(18): 4185β4192.
942:(9β10): 897β904.
787:(15): 3480β3493.
753:10.1002/bmb.21071
483:10.1115/1.4032188
108:The migration of
16:(Redirected from
3015:
2803:Crowd simulation
2780:Swarm algorithms
2751:Insect migration
2656:Animal migration
2648:Animal migration
2641:
2566:Mobbing behavior
2489:
2482:
2475:
2466:
2441:
2440:
2429:
2401:
2395:
2394:
2358:
2352:
2351:
2341:
2335:
2334:
2324:
2292:
2286:
2285:
2249:
2243:
2242:
2218:
2212:
2211:
2201:
2169:
2163:
2162:
2114:
2108:
2107:
2071:
2065:
2064:
2046:
2014:
2008:
2007:
1971:
1960:
1959:
1949:
1909:
1903:
1902:
1892:
1852:
1846:
1845:
1835:
1811:
1805:
1804:
1794:
1784:
1752:
1746:
1745:
1743:
1707:
1701:
1700:
1690:
1658:
1647:
1646:
1636:
1604:
1598:
1597:
1587:
1555:
1549:
1548:
1538:
1528:
1496:
1490:
1489:
1461:
1455:
1454:
1444:
1421:The EMBO Journal
1412:
1406:
1405:
1387:
1363:
1357:
1356:
1346:
1329:(6): 1265β1274.
1314:
1308:
1307:
1297:
1287:
1255:
1249:
1248:
1238:
1228:
1204:
1193:
1192:
1182:
1172:
1148:
1142:
1141:
1113:
1107:
1106:
1088:
1064:
1058:
1057:
1039:
1015:
1009:
1008:
998:
966:
960:
959:
931:
925:
924:
896:
890:
889:
887:
885:
866:
860:
859:
849:
821:
815:
814:
804:
781:The EMBO Journal
772:
766:
765:
755:
731:
725:
724:
714:
704:
672:
663:
659:
649:
609:
603:
602:
592:
560:
554:
553:
543:
511:
505:
504:
494:
462:
447:Protein dynamics
406:
401:
400:
383:inverse problems
63:vascular disease
51:chemical signals
47:immune responses
21:
3023:
3022:
3018:
3017:
3016:
3014:
3013:
3012:
2993:
2992:
2991:
2986:
2905:
2867:
2822:
2774:
2642:
2633:
2498:
2493:
2449:
2444:
2431:
2403:
2402:
2398:
2360:
2359:
2355:
2343:
2342:
2338:
2294:
2293:
2289:
2266:10.1038/nrm2522
2260:(11): 860β873.
2251:
2250:
2246:
2220:
2219:
2215:
2171:
2170:
2166:
2116:
2115:
2111:
2073:
2072:
2068:
2016:
2015:
2011:
1973:
1972:
1963:
1911:
1910:
1906:
1854:
1853:
1849:
1813:
1812:
1808:
1754:
1753:
1749:
1732:10.1119/1.10903
1709:
1708:
1704:
1660:
1659:
1650:
1606:
1605:
1601:
1557:
1556:
1552:
1498:
1497:
1493:
1463:
1462:
1458:
1414:
1413:
1409:
1365:
1364:
1360:
1316:
1315:
1311:
1257:
1256:
1252:
1206:
1205:
1196:
1150:
1149:
1145:
1115:
1114:
1110:
1066:
1065:
1061:
1017:
1016:
1012:
968:
967:
963:
933:
932:
928:
898:
897:
893:
883:
881:
868:
867:
863:
823:
822:
818:
774:
773:
769:
733:
732:
728:
687:(10): e110453.
674:
673:
666:
611:
610:
606:
562:
561:
557:
513:
512:
508:
464:
463:
459:
455:
432:Endocytic cycle
402:
395:
392:
379:
322:
313:
283:Reynolds number
263:
237:In the case of
228:endocytic cycle
224:
215:
202:
197:
189:
127:
125:Common features
106:
78:Reynolds number
67:tumor formation
28:
23:
22:
15:
12:
11:
5:
3021:
3019:
3011:
3010:
3005:
2995:
2994:
2988:
2987:
2985:
2984:
2979:
2974:
2969:
2964:
2962:Quorum sensing
2959:
2954:
2949:
2944:
2939:
2934:
2929:
2924:
2919:
2913:
2911:
2910:Related topics
2907:
2906:
2904:
2903:
2898:
2896:Swarm robotics
2893:
2888:
2883:
2877:
2875:
2873:Swarm robotics
2869:
2868:
2866:
2865:
2860:
2855:
2854:
2853:
2843:
2838:
2832:
2830:
2824:
2823:
2821:
2820:
2815:
2810:
2805:
2800:
2795:
2790:
2784:
2782:
2776:
2775:
2773:
2772:
2767:
2766:
2765:
2764:
2763:
2748:
2747:
2746:
2741:
2731:
2730:
2729:
2724:
2719:
2714:
2707:Fish migration
2704:
2702:Cell migration
2699:
2698:
2697:
2692:
2685:Bird migration
2682:
2681:
2680:
2678:coded wire tag
2675:
2674:
2673:
2663:
2652:
2650:
2644:
2643:
2636:
2634:
2632:
2631:
2626:
2621:
2616:
2615:
2614:
2604:
2603:
2602:
2597:
2587:
2586:
2585:
2575:
2574:
2573:
2571:feeding frenzy
2563:
2558:
2553:
2552:
2551:
2541:
2540:
2539:
2534:
2524:
2519:
2514:
2508:
2506:
2500:
2499:
2494:
2492:
2491:
2484:
2477:
2469:
2463:
2462:
2456:
2448:
2447:External links
2445:
2443:
2442:
2396:
2369:(2): 169β179.
2353:
2336:
2307:(6): 721β729.
2287:
2244:
2213:
2164:
2109:
2066:
2009:
1982:(3): 658β682.
1961:
1904:
1847:
1806:
1747:
1702:
1673:(1): 9β22.e4.
1648:
1599:
1550:
1491:
1456:
1427:(2): 405β410.
1407:
1378:(4): 601β606.
1358:
1309:
1270:(2): 454β458.
1250:
1194:
1143:
1108:
1079:(4): 453β465.
1059:
1030:(3): 371β379.
1010:
981:(2): 597β602.
961:
926:
907:(2): 389β398.
891:
861:
816:
767:
746:(6): 475β482.
726:
664:
604:
575:(5): 608β623.
555:
506:
456:
454:
451:
450:
449:
444:
439:
434:
429:
424:
419:
414:
408:
407:
404:Biology portal
391:
388:
378:
375:
330:surface charge
321:
318:
312:
309:
262:
259:
223:
220:
214:
211:
201:
198:
196:
193:
188:
185:
146:
145:
142:
126:
123:
121:cell forward.
110:cultured cells
105:
102:
31:Cell migration
26:
24:
18:Cell Migration
14:
13:
10:
9:
6:
4:
3:
2:
3020:
3009:
3008:Cell movement
3006:
3004:
3001:
3000:
2998:
2983:
2980:
2978:
2975:
2973:
2970:
2968:
2965:
2963:
2960:
2958:
2955:
2953:
2950:
2948:
2945:
2943:
2940:
2938:
2935:
2933:
2930:
2928:
2925:
2923:
2920:
2918:
2915:
2914:
2912:
2908:
2902:
2899:
2897:
2894:
2892:
2889:
2887:
2884:
2882:
2879:
2878:
2876:
2874:
2870:
2864:
2861:
2859:
2856:
2852:
2849:
2848:
2847:
2844:
2842:
2839:
2837:
2836:Active matter
2834:
2833:
2831:
2829:
2825:
2819:
2816:
2814:
2811:
2809:
2806:
2804:
2801:
2799:
2796:
2794:
2791:
2789:
2786:
2785:
2783:
2781:
2777:
2771:
2768:
2762:
2759:
2758:
2757:
2754:
2753:
2752:
2749:
2745:
2742:
2740:
2737:
2736:
2735:
2732:
2728:
2725:
2723:
2720:
2718:
2715:
2713:
2712:diel vertical
2710:
2709:
2708:
2705:
2703:
2700:
2696:
2693:
2691:
2688:
2687:
2686:
2683:
2679:
2676:
2672:
2669:
2668:
2667:
2664:
2662:
2659:
2658:
2657:
2654:
2653:
2651:
2649:
2645:
2640:
2630:
2627:
2625:
2622:
2620:
2617:
2613:
2610:
2609:
2608:
2605:
2601:
2598:
2596:
2593:
2592:
2591:
2588:
2584:
2581:
2580:
2579:
2576:
2572:
2569:
2568:
2567:
2564:
2562:
2559:
2557:
2554:
2550:
2549:herd behavior
2547:
2546:
2545:
2542:
2538:
2535:
2533:
2530:
2529:
2528:
2525:
2523:
2520:
2518:
2515:
2513:
2510:
2509:
2507:
2505:
2501:
2497:
2490:
2485:
2483:
2478:
2476:
2471:
2470:
2467:
2460:
2457:
2454:
2451:
2450:
2446:
2438:
2434:
2427:
2423:
2419:
2415:
2412:(3): 658β82.
2411:
2407:
2400:
2397:
2392:
2388:
2384:
2380:
2376:
2372:
2368:
2364:
2357:
2354:
2349:
2348:
2340:
2337:
2332:
2328:
2323:
2318:
2314:
2310:
2306:
2302:
2298:
2291:
2288:
2283:
2279:
2275:
2271:
2267:
2263:
2259:
2255:
2248:
2245:
2240:
2236:
2232:
2228:
2224:
2217:
2214:
2209:
2205:
2200:
2195:
2191:
2187:
2183:
2179:
2175:
2168:
2165:
2160:
2156:
2152:
2148:
2144:
2140:
2136:
2132:
2128:
2124:
2120:
2113:
2110:
2105:
2101:
2097:
2093:
2089:
2085:
2081:
2077:
2070:
2067:
2062:
2058:
2054:
2050:
2045:
2040:
2036:
2032:
2028:
2024:
2020:
2013:
2010:
2005:
2001:
1997:
1993:
1989:
1985:
1981:
1977:
1970:
1968:
1966:
1962:
1957:
1953:
1948:
1943:
1939:
1935:
1931:
1927:
1923:
1919:
1915:
1908:
1905:
1900:
1896:
1891:
1886:
1882:
1878:
1874:
1870:
1866:
1862:
1858:
1851:
1848:
1843:
1839:
1834:
1829:
1825:
1821:
1817:
1810:
1807:
1802:
1798:
1793:
1788:
1783:
1778:
1774:
1770:
1766:
1762:
1758:
1751:
1748:
1742:
1737:
1733:
1729:
1725:
1721:
1717:
1713:
1706:
1703:
1698:
1694:
1689:
1684:
1680:
1676:
1672:
1668:
1664:
1657:
1655:
1653:
1649:
1644:
1640:
1635:
1630:
1626:
1622:
1618:
1614:
1610:
1603:
1600:
1595:
1591:
1586:
1581:
1577:
1573:
1569:
1565:
1561:
1554:
1551:
1546:
1542:
1537:
1532:
1527:
1522:
1518:
1514:
1510:
1506:
1502:
1495:
1492:
1487:
1483:
1479:
1475:
1471:
1467:
1460:
1457:
1452:
1448:
1443:
1438:
1434:
1430:
1426:
1422:
1418:
1411:
1408:
1403:
1399:
1395:
1391:
1386:
1381:
1377:
1373:
1369:
1362:
1359:
1354:
1350:
1345:
1340:
1336:
1332:
1328:
1324:
1320:
1313:
1310:
1305:
1301:
1296:
1291:
1286:
1281:
1277:
1273:
1269:
1265:
1261:
1254:
1251:
1246:
1242:
1237:
1232:
1227:
1222:
1218:
1214:
1210:
1203:
1201:
1199:
1195:
1190:
1186:
1181:
1176:
1171:
1166:
1162:
1158:
1154:
1147:
1144:
1139:
1135:
1131:
1127:
1123:
1119:
1112:
1109:
1104:
1100:
1096:
1092:
1087:
1082:
1078:
1074:
1070:
1063:
1060:
1055:
1051:
1047:
1043:
1038:
1033:
1029:
1025:
1021:
1014:
1011:
1006:
1002:
997:
992:
988:
984:
980:
976:
972:
965:
962:
957:
953:
949:
945:
941:
937:
930:
927:
922:
918:
914:
910:
906:
902:
895:
892:
879:
875:
871:
865:
862:
857:
853:
848:
843:
839:
835:
831:
827:
820:
817:
812:
808:
803:
798:
794:
790:
786:
782:
778:
771:
768:
763:
759:
754:
749:
745:
741:
737:
730:
727:
722:
718:
713:
708:
703:
698:
694:
690:
686:
682:
678:
671:
669:
665:
662:
657:
653:
648:
643:
639:
635:
631:
627:
623:
619:
615:
608:
605:
600:
596:
591:
586:
582:
578:
574:
570:
566:
559:
556:
551:
547:
542:
537:
533:
529:
525:
521:
517:
510:
507:
502:
498:
493:
488:
484:
480:
477:(2): 021004.
476:
472:
468:
461:
458:
452:
448:
445:
443:
440:
438:
435:
433:
430:
428:
425:
423:
420:
418:
415:
413:
412:Cap formation
410:
409:
405:
399:
394:
389:
387:
384:
376:
374:
370:
368:
363:
358:
353:
351:
347:
343:
339:
335:
331:
327:
319:
317:
310:
304:
300:
297:
293:
289:
286:both able to
284:
280:
279:E. M. Purcell
276:
272:
268:
267:Dictyostelium
260:
254:
250:
247:
243:
241:
240:Dictyostelium
235:
233:
229:
221:
219:
213:Trailing edge
212:
210:
207:
199:
194:
192:
186:
180:
172:
168:
166:
162:
158:
153:
151:
143:
140:
139:
138:
136:
132:
131:spermatozooic
124:
122:
119:
115:
111:
103:
101:
99:
95:
91:
87:
83:
79:
74:
72:
68:
64:
60:
56:
52:
48:
44:
43:wound healing
40:
36:
32:
19:
2917:Allee effect
2891:Nanorobotics
2881:Ant robotics
2858:Vicsek model
2701:
2437:ScienceDaily
2436:
2409:
2405:
2399:
2366:
2362:
2356:
2346:
2339:
2304:
2300:
2290:
2257:
2253:
2247:
2222:
2216:
2181:
2177:
2167:
2126:
2122:
2112:
2079:
2075:
2069:
2026:
2022:
2012:
1979:
1975:
1921:
1917:
1907:
1867:(1): 12970.
1864:
1860:
1850:
1823:
1819:
1809:
1764:
1760:
1750:
1715:
1711:
1705:
1670:
1666:
1616:
1612:
1602:
1567:
1563:
1553:
1508:
1504:
1494:
1469:
1465:
1459:
1424:
1420:
1410:
1375:
1371:
1361:
1326:
1322:
1312:
1267:
1263:
1253:
1216:
1212:
1160:
1156:
1146:
1121:
1117:
1111:
1076:
1072:
1062:
1027:
1023:
1013:
978:
974:
964:
939:
935:
929:
904:
900:
894:
882:. Retrieved
878:the original
873:
864:
838:10.3791/3589
829:
819:
784:
780:
770:
743:
739:
729:
684:
680:
624:(1): 1β112.
621:
617:
607:
572:
568:
558:
523:
519:
509:
474:
470:
460:
380:
371:
357:microtubules
354:
334:cell surface
323:
314:
295:
292:optogenetics
266:
264:
239:
236:
225:
216:
203:
200:Leading edge
190:
154:
147:
128:
107:
94:cytoskeleton
75:
30:
29:
2937:Eusociality
2886:Microbotics
2756:butterflies
2727:sardine run
2661:altitudinal
2583:pack hunter
1741:2433/226838
1718:(3): 3β11.
1511:(8): e724.
1466:Development
526:: 265β289.
442:Neurophilic
275:macrophages
271:neutrophils
2997:Categories
2851:clustering
2744:philopatry
2722:salmon run
2717:Lessepsian
1826:(1): 1β3.
453:References
417:Chemotaxis
346:Rho GTPase
161:cyclic AMP
135:locomotion
114:microscopy
90:Eukaryotic
71:metastasis
2972:Stigmergy
2952:Mutualism
2612:bait ball
2061:248990694
1124:: 65β95.
427:Durotaxis
367:receptors
269:amoebae,
232:integrins
2901:Symbrion
2863:BIO-LGCA
2666:tracking
2595:ant mill
2537:sort sol
2532:flocking
2496:Swarming
2426:20878250
2391:16997326
2331:22665521
2282:19500145
2274:18946475
2239:38951708
2225:: 1β15.
2208:37220748
2199:10524337
2159:16029926
2151:14657486
2104:10221901
2053:36202973
2044:10029748
2004:37036941
1996:20878250
1956:30337409
1899:29021607
1842:29974859
1801:20534502
1697:29937389
1643:10588667
1594:20807800
1545:17684569
1505:PLOS ONE
1486:12183371
1402:14776455
1245:23135278
1189:20696757
1138:18573073
1095:12600310
956:16962888
884:24 March
856:22215133
811:16900100
762:28627731
721:25310593
681:PLOS ONE
656:24748680
599:32877650
550:20192768
501:26639083
390:See also
362:vesicles
326:polarity
288:chemotax
150:antibody
98:blebbing
82:flagella
2761:monarch
2690:flyways
2671:history
2522:Droving
2371:Bibcode
2322:3373410
2131:Bibcode
2123:Science
2084:Bibcode
2076:Science
1947:6218007
1926:Bibcode
1918:Science
1890:5636814
1869:Bibcode
1792:2895083
1769:Bibcode
1720:Bibcode
1688:6048972
1585:2939799
1536:1933600
1513:Bibcode
1451:1531629
1394:8929529
1353:7515888
1344:2290921
1304:6300844
1272:Bibcode
1236:3527923
1180:2952225
1103:6887118
1046:8608590
1005:4040521
996:2113673
921:5531377
847:3369670
802:1538568
712:4195729
689:Bibcode
647:3985726
626:Bibcode
590:7505206
541:4364543
492:4844084
242:amoebae
165:haploid
2734:Homing
2556:Locust
2424:
2389:
2329:
2319:
2280:
2272:
2237:
2206:
2196:
2157:
2149:
2102:
2059:
2051:
2041:
2002:
1994:
1954:
1944:
1897:
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1840:
1799:
1789:
1695:
1685:
1641:
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1592:
1582:
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1449:
1442:556468
1439:
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1341:
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1292:
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1054:982415
1052:
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799:
760:
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709:
661:online
654:
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587:
548:
538:
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489:
2798:Boids
2739:natal
2527:Flock
2278:S2CID
2155:S2CID
2057:S2CID
2000:S2CID
1634:25767
1398:S2CID
1099:S2CID
1050:S2CID
342:CDC42
206:actin
86:cilia
2578:Pack
2544:Herd
2422:PMID
2387:PMID
2327:PMID
2270:PMID
2235:PMID
2204:PMID
2147:PMID
2100:PMID
2049:PMID
1992:PMID
1952:PMID
1895:PMID
1838:PMID
1797:PMID
1693:PMID
1639:PMID
1590:PMID
1541:PMID
1482:PMID
1447:PMID
1390:PMID
1372:Cell
1349:PMID
1300:PMID
1241:PMID
1185:PMID
1134:PMID
1091:PMID
1073:Cell
1042:PMID
1024:Cell
1001:PMID
952:PMID
917:PMID
886:2013
852:PMID
807:PMID
758:PMID
717:PMID
652:PMID
595:PMID
546:PMID
497:PMID
350:PTEN
348:and
338:PIP3
69:and
53:and
45:and
2414:doi
2379:doi
2367:244
2317:PMC
2309:doi
2305:197
2262:doi
2227:doi
2194:PMC
2186:doi
2139:doi
2127:302
2092:doi
2080:284
2039:PMC
2031:doi
1984:doi
1942:PMC
1934:doi
1922:362
1885:PMC
1877:doi
1828:doi
1787:PMC
1777:doi
1765:107
1736:hdl
1728:doi
1683:PMC
1675:doi
1629:PMC
1621:doi
1580:PMC
1572:doi
1568:123
1531:PMC
1521:doi
1474:doi
1470:129
1437:PMC
1429:doi
1380:doi
1339:PMC
1331:doi
1327:125
1290:PMC
1280:doi
1231:PMC
1221:doi
1217:287
1175:PMC
1165:doi
1161:285
1126:doi
1081:doi
1077:112
1032:doi
991:PMC
983:doi
979:101
944:doi
909:doi
842:PMC
834:doi
797:PMC
789:doi
748:doi
707:PMC
697:doi
642:PMC
634:doi
585:PMC
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536:PMC
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