Meta-review: trends and predictions

A curated, semi-automated review of thousands of bleb papers: how they were found, read and counted, what the counts show, and what past and present reviews expected of the field.

This page is a meta-review: a review of the bleb literature as a whole rather than of any single question. Searches of databases and of the older literature, followed by sorting under written rules that we tested by hand, produced nine curated paper lists holding 5,175 dated papers from 1856 to 2026, and structured reading notes cover about 5,400 publications. We use this material to count how bleb research has grown and shifted, and to compare what reviews expected of the field with what happened. Counts measure activity, not importance; which work changed ideas is the subject of A long history of blebbing.

A map of the field

The map links the main ideas on this site: the molecules that set the mechanics of the cell surface, the forces and materials involved, the life of a single bleb, the kinds of blebs, and the cells and conditions in which they appear. Each arrow states a relation that the linked pages describe and cite; dashed lines join related ideas without implying cause.

Swipe sideways to see the whole map.

Concept map: how the main ideas about blebs connectRho–ROCKmyosin IIblebbistatinactin turnoveractin cortexcortical tensionLaplace's lawintracellularpressureporoelasticityERM proteinsmembrane–cortexattachmentreconstitutioninverse blebblebinitiation:detachmentor ruptureexpansionnew cortexassemblesretractioncortex scarcortical flowpercolationcytokineplaststable blebleader blebconfinementcircus blebfrictionbleb-basedmigrationtransient blebmesenchymal–amoeboidtransitionlamellipodiumseptinsignaling hubapoptotic blebapoptotic bodylarge oncosomenuclear blebactivatesinhibitscontractsrenewsgeneratesraisesrelatesrelatesdelays its spreadprovideopposesdrivesdrivesif the outside pressure is higherreproduces both routesthenstops whenthenleaves the old cortexstarts bya full cyclere-forms on one side only (proposed)keeps the tip free of cortexorganizes (proposed)with low adhesion, inducesclosely relatedcan detach asfast, under confinementwhen polarizedtransmits force inswitches cells tothe alternative to blebs at the frontrepeated blebbing gathersin dying cellsmay pinch off assaid to pinch off asnot the same: nuclear envelope

Hover over or tap a concept to see what it connects to; select it to read its definition and find the pages that treat it. The buttons hide or show each kind of concept.

All connections as a list
  • Rho–ROCK → myosin II: activates
  • blebbistatin → myosin II: inhibits
  • myosin II → actin cortex: contracts
  • actin turnover → actin cortex: renews
  • actin cortex → cortical tension: generates
  • cortical tension → intracellular pressure: raises
  • Laplace's law – cortical tension: relates
  • Laplace's law – intracellular pressure: relates
  • poroelasticity → intracellular pressure: delays its spread
  • ERM proteins → membrane–cortex attachment: provide
  • membrane–cortex attachment → initiation: detachment or rupture: opposes
  • intracellular pressure → expansion: drives
  • intracellular pressure → initiation: detachment or rupture: drives
  • intracellular pressure → inverse bleb: if the outside pressure is higher
  • reconstitution → initiation: detachment or rupture: reproduces both routes
  • initiation: detachment or rupture → expansion: then
  • expansion → new cortex assembles: stops when
  • new cortex assembles → retraction: then
  • initiation: detachment or rupture → cortex scar: leaves the old cortex
  • bleb → initiation: detachment or rupture: starts by
  • retraction → transient bleb: a full cycle
  • new cortex assembles → circus bleb: re-forms on one side only (proposed)
  • cortical flow → stable bleb: keeps the tip free of cortex
  • percolation → cortical flow: organizes (proposed)
  • confinement → stable bleb: with low adhesion, induces
  • stable bleb – leader bleb: closely related
  • stable bleb → cytokineplast: can detach as
  • stable bleb → bleb-based migration: fast, under confinement
  • transient bleb → bleb-based migration: when polarized
  • friction → bleb-based migration: transmits force in
  • mesenchymal–amoeboid transition → bleb-based migration: switches cells to
  • lamellipodium – bleb-based migration: the alternative to blebs at the front
  • transient bleb → septin signaling hub: repeated blebbing gathers
  • bleb → apoptotic bleb: in dying cells
  • apoptotic bleb → apoptotic body: may pinch off as
  • bleb → large oncosome: said to pinch off as
  • bleb – nuclear bleb: not the same: nuclear envelope

How the review was built

Searching, sorting, note-taking, tagging and counting were automated, carried out by code and by language models working under written rules. On 26 September 2026 we searched Europe PMC and PubMed for "bleb" in all its forms and for four older terms, zeiosis, zeiotic, hyaline cap and circus movement, in titles, abstracts and the main sections of open-access articles, with a separate search for amoeboid movement. Merging records that shared a DOI, PubMed or PubMed Central identifier left 57,430 works. Named rules flagged common false hits such as glaucoma, lung and skin "blebs" and bacterial membrane vesicles, and after 1950 only English-language work was kept.

A word search cannot tell a study of blebs from a passing mention, so a language model sorted the candidates against a written codebook. A paper counts as a bleb paper if it studies blebs of living cells or reports them in its results, but not if blebbing serves only as a sign of death or toxicity. The model sorted all 7,207 papers with a bleb word in the title or abstract and 3,641 of the 16,503 that used the word only inside the article, after fixed rules had set aside those without open full text and those that used the word once, outside the main sections, or only about cell death or vesicles. Work from 1800 to 1950, before the words for the phenomenon settled, came from a separate search of scanned journals, databases and historical reference lists, which gave 4,566 records sorted by title.

Each listed paper then received a structured note, written by a language model to a fixed protocol that records the system, the observations and numbers, the authors' interpretation, the earlier work they credit, and their predictions and open questions. Depth followed relevance and access: 704 papers were read deeply with every figure, 1,571 at standard depth and the rest briefly, and only legally open texts and PDFs from our own collection were read. A language model assigned topic tags from titles and abstracts, and a script computed every count on this page from the lists, notes and tags as they stood on 30 September 2026.

The judgment that shaped the lists was ours. We decided by hand what counts as a bleb paper: we revised the codebook after each of two audits of 100 sorting decisions, and the bulk-sorting rules, tested on the second audit before a last revision, agreed with our answers on 95 of 100. A set of 224 publications from our own reference library served as a seed and as a test of completeness. Before sorting, the searches found 111 of the 117 items in it that use a bleb word, missing only works the databases do not index, and all 106 bleb papers cited by five reviews from other groups. A citation check of the 200 works that at least five kept papers cite, but that the searches had missed or the rules had cut, recovered nine bleb papers, among them Paluch et al. (2005).

What the counts show

The bleb list holds 2,426 counted papers once two retracted papers are set aside. It grew from about 23 papers a year in 1975–2004 to 106 a year in 2015–2024, with a peak of 141 in 2021, so half of all bleb papers appeared in 2015 or later. The nuclear-bleb list, which concerns a different membrane, grew faster still, from 54 papers in 2005–2014 to 245 in 2015–2026.

Papers per publication year in the paper lists, with the three borderline lists shown together; each paper is counted once, in the first list of the legend that holds it. The rise in the mid-1970s is partly a database artifact, and 2026 is…
Papers per publication year in the paper lists, with the three borderline lists shown together; each paper is counted once, in the first list of the legend that holds it. The rise in the mid-1970s is partly a database artifact, and 2026 is a partial year. Source: blebology.com, counts of 30 September 2026.

What the papers are about shifted twice. Cell death accounted for 10% of papers in 1975–1989, 30% in 1990–2004 and 43% in 2000–2004, the highest share of any setting in any period since 1975; since papers that use blebbing merely as a marker of death were excluded, these papers study the blebbing of dying cells. Migration then took over, rising from about 8% of papers before 2005 to 18% in 2005–2014 and 21% in 2015–2026, the most common of the five settings in every five-year period since 2015, while cell death fell back to 16%. Development fell from 16% in 1975–1989 to about 3% in 2000–2004 and has risen again, to 12% in 2025–2026, and reconstitution in cell-free or vesicle systems rose from 1% or less before 2005 to 4% in 2015–2026.

Share of papers in five settings per five-year period, counted in the bleb, dying-cell and reconstituted lists; a paper can count in several settings, and on phones the figure omits the number of papers per period. Note the cell-death peak…
Share of papers in five settings per five-year period, counted in the bleb, dying-cell and reconstituted lists; a paper can count in several settings, and on phones the figure omits the number of papers per period. Note the cell-death peak in 2000–2004 and the lead of migration from 2015. Source: blebology.com, counts of 30 September 2026.

The systems and tools changed with the questions. Among bleb papers that name an organism, human cells rose from 27% in 1975–1989 to 55% in 2015–2026, mouse and rat cells fell from 37% to 17%, and fish, flies, worms and amoebae together grew from about 3% to about 17%. Among papers read in full, over the same periods, electron microscopy fell from 73% to 22%, while fluorescence imaging rose from 31% to 85%, genetic perturbation from 6% to more than half, engineered environments such as gels and microchannels from 12% to 39%, and modeling from 4% to 21%. The early method shares rest on only 81 papers read in full, so they are rough.

Organism studied in bleb papers, as a share of papers that state one, per five-year period, with years before 1975 pooled; reviews and theory are left out. Look at the rise of human cells and the fall of rodent cells.
Organism studied in bleb papers, as a share of papers that state one, per five-year period, with years before 1975 pooled; reviews and theory are left out. Look at the rise of human cells and the fall of rodent cells. Source: blebology.com, counts of 30 September 2026.
Percentage of papers read in full, reviews excluded, that report each kind of method, by period; n is given under each column. Electron microscopy gives way to fluorescence imaging and genetic perturbation.
Percentage of papers read in full, reviews excluded, that report each kind of method, by period; n is given under each column. Electron microscopy gives way to fluorescence imaging and genetic perturbation. Source: blebology.com, counts of 30 September 2026.

The vocabulary settled as the field grew. "Bleb" in some form occurs in 80% of the 86 full texts searched from 1975–1989 and in 91% of the 1,547 from 2015–2026, while "zeiosis" fell from 20% to 0.1%. Newer words name modes and products rather than shapes: phrases for bleb-based migration occur in 13% of the full texts from 2015–2026 and "extracellular vesicle" in 15%, and leader bleb, inverse bleb and blebbisome first appear in our notes between 2015 and 2025.

Words for blebs, amoeboid movement and related structures: papers using each word per five-year period (dot size), with the author and year of the earliest use found (label); on phones the figure shows only the 15 words used in at least 50…
Words for blebs, amoeboid movement and related structures: papers using each word per five-year period (dot size), with the author and year of the earliest use found (label); on phones the figure shows only the 15 words used in at least 50 papers. A first use is the earliest in our notes, not necessarily the coinage; the Glossary gives the context of each. Source: blebology.com, counts of 30 September 2026.

How far the counts can be trusted

The searches define the vocabulary. Because the lists were found with bleb words and four older terms, the dominance of "bleb" is partly built in, and papers that say only "blister", "bubble" or "lobopodium" enter mainly through full text or the citation check. Database coverage also changes: before about 1975 PubMed records rarely carry abstracts, so the jump of the bleb list from 14 papers in 1970–1974 to 124 in 1975–1979 is probably partly an artifact. The historical list was sorted from titles alone and is not comparable with the later lists.

Access is the largest source of bias. Searching inside articles reached only open-access papers, about a third of the bleb papers were found only that way, and the 5,303 papers without open full text were not examined at all. The rules that thinned the full-text hits may also have lost about one relevant paper in ten, a rough estimate from a test of ten papers. Only 15% of listed papers from 1975–2004 were read in full, against 71% of those from 2010–2024, and 70% of the records from before 1950 carry only a title and journal, so trends for the older periods rest mostly on abstracts. Growth in open access should not be mistaken for growth of the field.

Reading depth by five-year period. Top, papers listed; bottom, the share read in full (text or scan), from the abstract only, or from the bibliographic record only. Most papers from 1975–2004 were read from abstracts, and most records…
Reading depth by five-year period. Top, papers listed; bottom, the share read in full (text or scan), from the abstract only, or from the bibliographic record only. Most papers from 1975–2004 were read from abstracts, and most records before 1950 carry only a title and journal. Source: blebology.com, counts of 30 September 2026.

Sorting has an expected error of about 5%, from the audit of 100 decisions, while the topic tags were only spot-checked. The tags follow the categories of one review (García-Arcos et al., 2024a ◆), so we checked them against how papers by other groups distribute. Between 55% and 74% of bleb papers in each period since 1975 carry only the code "function or phenotype": they report blebs, or what blebs do, without studying how blebs form. The counts are therefore a map of activity rather than of understanding.

What reviews up to 2018 predicted, and what happened

To judge how far reviews can guide expectations, we went back to 147 reviews, perspectives, books and theses dated 1764 to 2018, extracted 1,471 predictions and open questions, and scored each against later primary papers in the reading notes. Language-model readers did the scoring under written rules: only primary papers published after the source counted, and a later review could mark a point as still open but never as answered. About half of the items, 721, concerned matters the lists were not built to cover. Of the 750 that could be tested, 17% were realized, 43% partly realized, 38% not yet and 1.5% contradicted. For sources from 2005–2018, the first full answer came a median of 5 years after the review and the first partial answer a median of 4 years. "Not yet" means not found in these lists, and because the lists were partly assembled around the papers that answered the old questions, the record is flattered.

The clearest success concerned migration without adhesion. Charras and Paluch (2008) proposed that cells squeezed between surfaces could bleb forward without specific adhesion, whereas Lämmermann and Sixt (2009) argued that blebbing tumor cells draw their traction from residual adhesive protrusions. In 2015, confined cells of many types migrated fast on non-adhesive surfaces, driven by rearward cortical flow that grips the walls by friction (Bergert et al., 2015; Liu et al., 2015; Ruprecht et al., 2015). The switch is established for confined cells in vitro (M20), and 2015 alone brought the first answer to 47 of the old items, 32 of them through four papers on confined cells.

Other forecasts came true in part, often because a general question received a different answer in each system. Charras (2008) asked, for example, whether lipid flows into growing blebs. It did not in zebrafish germ cells, where invaginations of the surface unfolded instead (Goudarzi et al., 2017), but it flowed from base to tip in the stable leader blebs of melanoma cells (Jha et al., 2026). The answer depends on the kind of bleb, which the question had not separated.

Some questions remain open after nearly two decades. Reviews of 2006–2008 asked whether blebs in migrating cells start where the membrane detaches from an intact cortex, the actin layer beneath it, or where the cortex tears (rupture versus detachment). Reconstituted cortices in liposomes show that both routes are possible (Miyazaki et al., 2026), and in single examples from nucleus-free fragments of L929 cells the cortex tore before the bulge grew (Paluch et al., 2005). In Dictyostelium, however, one study saw the old cortex persist intact after each bleb (Zatulovskiy et al., 2014) and another found narrow gaps at inferred nucleation sites (Asante-Asamani et al., 2022). In our reading, a narrow gap is easy to miss in a survey, and the targeted analysis is correlative. Whether migrating vertebrate cells start blebs by rupture is an open question (M15).

Some kinds of forecast fared better than others. Calls for a new measurement, tool or experimental system were fully realized in 28% of cases, against 17% for other statements, probably because they usually describe work already under way. Statements about physical quantities such as tension, pressure or friction did better than those naming only molecules (26% against 18%), forecasts set in a living organism or tissue did worse (12% against 22% where no setting was named), and therapy, biomarker and clinical forecasts fared worst, with 4% realized and 65% not yet. Five of the 11 contradictions came from claims that one mechanism was universal or exclusive, such as the proposal that an adhesion–RhoA–ROCK–myosin II axis controls all migration in three dimensions (Petrie and Yamada, 2012), whose adhesion arm was contradicted by adhesion-free migration in confinement (Bergert et al., 2015). Forecasters also often answered their own questions: for 28 of 105 realized modern items, the first qualifying paper shared an author surname with the review.

Where current reviews expect the field to go

To see where the field is heading, we read the 323 reviews, perspectives, commentaries, books and theses from 2019–2026 in the lists, 31 of them only as abstracts and 8 from the record alone. Most mention blebs only in passing: about 110 are reviews of extracellular vesicles that repeat the textbook link between blebbing and vesicle release, most of the 26 on the nucleus use "bleb" for herniations of the nuclear envelope, about 50 are off-topic, and only 17 are centered on blebs. On bleb nucleation, membrane tension and theory, primary papers are ahead of the reviews. We grouped the directions that reviewers name into six themes and set each against the record of similar forecasts in the backward check, which describes kinds of forecasts, not the odds for any one.

Whether blebs are needed in living tissue. The direction named most often, by about fifteen reviews and commentaries, is to show that blebs are required in tissues, not merely present (Schick and Raz, 2022; Caruso and Logue, 2024; García-Arcos et al., 2024a ◆). The difficulty is that inhibiting myosin or ROCK, the usual way to stop blebs, also lowers the contractility this migration needs, so blebs could be a by-product (Schick and Raz, 2022). Perturbations aimed at the bleb itself are beginning to appear: chick germ cells carrying a dominant-negative form of the calcium channel Orai1 rarely blebbed and left the blood vessels about 40% less efficiently (4 embryos; Morita et al., 2026). Mouse primordial germ cells, in contrast, migrate with actin-rich protrusions and bleb only when actin polymerization is blocked (Goodwin et al., 2026). Forecasts set in a living organism were realized in only 12% of cases.

The physics of a single bleb. Roughly six to eleven reviews each ask how a bleb starts, where its extra membrane comes from, how water and pressure move as it grows, and whether the connectivity of the cortex controls it (Ikenouchi and Aoki, 2022; Fujii and Ikenouchi, 2024). Water flow and pressure are still inferred rather than measured: an osmotic model of bleb growth needs about twice the protein concentration difference that was measured (Fujii et al., 2026). Cortex connectivity as a control variable rests largely on one cell line (García-Arcos et al., 2024b ◆) and one liposome study. In the backward check, such mechanistic questions took longest to answer, a median of six to seven years, and the answers usually held in one system.

Rebuilding blebs from purified parts. A smaller group expects blebbing to be rebuilt from purified components, first as single blebs, then as blebs that retract and form again, and eventually as movement. Only one recent review discusses it (Lopes Dos Santos and Campillo, 2022). Vesicles with an actomyosin shell formed static blebs where the shell was less firmly anchored to the membrane (Loiseau et al., 2016), and laser cuts in a model cortex triggered blebs in at most 13% of liposomes (Sakamoto and Murrell, 2024), but none of these blebs has yet retracted by rebuilding its cortex (M15). Calls for a method, measurement or reconstituted system have the best record in the backward check, fully realized in 28–31% of cases.

Blebs as signaling sites, vesicles and fragments. About fifteen reviews and commentaries treat blebs as compartments that concentrate signals or leave the cell (Reichman-Fried and Raz, 2023; Di Vizio et al., 2023; García-Arcos et al., 2024a ◆), but the evidence is narrower than the attention. The view of blebs as signaling hubs rests mainly on septin-organized survival signaling in detached melanoma cells in vitro (Weems et al., 2023) and on polarized signaling modules in leader blebs (Jha et al., 2026). The expectation that blebs give rise to motile fragments in the body rests on stable blebs detaching in one confined cell line (García-Arcos et al., 2024b ◆) and on lung fragments whose bleb origin was not tested (Headley et al., 2016); here the forecast and the main evidence come from the same group. Blebbisomes, vesicles that keep blebbing for days, form in a retraction rather than from a bleb (Jeppesen et al., 2025). Of past forecasts of this kind, none of 43 testable items on signaling machinery and 6% of 193 on vesicles were fully realized.

Treating amoeboid cancer cells. Cancer reviews agree more closely than any other group. About ten expect amoeboid migration, the rounded, contractile and bleb-rich mode of some invading cancer cells, to become a drug target, because amoeboid cells gather at invasive fronts and depend on Rho–ROCK–myosin II contractility (Graziani et al., 2022; Jia et al., 2024; Ghose and Sanz-Moreno, 2026). ROCK inhibitors combined with drugs that block escape into other modes should then slow dissemination, although even Jia et al. (2024) judge clinical translation to be distant. Forecasts of this kind have the weakest record. Systemic ROCK inhibition reduced growth, invasion and lung metastasis in a mouse melanoma model (5 control and 4 treated mice; Rodriguez-Hernandez et al., 2020), but AT13148, a ROCK-targeting drug tested in a phase I cancer trial, was closed for adverse effects and lack of response (Graziani et al., 2022).

New ways to classify migration, and the origin of blebbing. About eight reviews expect migration to be classified by how force is generated and transmitted rather than by protrusion shape, because one cell can combine blebs with lamellipodia and the modes may form a continuum (Bodor et al., 2020; Alonso-Matilla et al., 2025; Duggan and Petrie, 2025). Choanoflagellates, the closest living relatives of animals, bleb when confined (Brunet et al., 2021), and one review proposes that migration without adhesions is older than migration driven by focal adhesions (Bodor et al., 2020). Past forecasts here held when they predicted variety, with new modes found within two to five years, and failed when they claimed generality. Whether blebs are older than lamellipodia, asked by Charras and Paluch (2008), is untested, so this expectation is an agenda rather than a forecast.

In our reading, the two checks together suggest how to read today's expectations. Partial answers tend to arrive within about four years, usually in one cell type or species. Calls for methods and measurements deserve more weight than forecasts of clinical benefit, and words such as "universal" or "the main determinant" deserve caution. The lists behind these counts can be downloaded from Paper lists.

References (38)
  • Alonso-Matilla R, Provenzano PP, Odde DJ (2025) Physical principles and mechanisms of cell migration. npj Biol Phys Mech 2:2. doi:10.1038/s44341-024-00008-w
  • Asante-Asamani E, Grange D, Rawal D et al. (2022) A role for myosin II clusters and membrane energy in cortex rupture for Dictyostelium discoideum. PLoS ONE e0265380. doi:10.1371/journal.pone.0265380
  • Bergert M, Erzberger A, Desai RA et al. (2015) Force transmission during adhesion-independent migration. Nat Cell Biol 17:524–529. doi:10.1038/ncb3134
  • Bodor DL, Pönisch W, Endres RG et al. (2020) Of cell shapes and motion: the physical basis of animal cell migration. Dev Cell 52:550–562. doi:10.1016/j.devcel.2020.02.013
  • Brunet T, Albert M, Roman W et al. (2021) A flagellate-to-amoeboid switch in the closest living relatives of animals. eLife 10:e61037. doi:10.7554/elife.61037
  • Caruso AP, Logue JS (2024) The biophysics of cell motility through mechanochemically challenging environments. Curr Opin Cell Biol 90:102404. doi:10.1016/j.ceb.2024.102404
  • Charras GT (2008) A short history of blebbing. J Microsc 231:466–478. doi:10.1111/j.1365-2818.2008.02059.x
  • Charras G, Paluch E (2008) Blebs lead the way: how to migrate without lamellipodia. Nat Rev Mol Cell Biol 9:730–736. doi:10.1038/nrm2453
  • Di Vizio D, Schoppet M, Weeraratna A et al. (2023) Blebs and former blebs: from surface protrusions to extracellular vesicles in cancer signalling, anoikis resistance and beyond. J Extracell Biol 2:e112. doi:10.1002/jex2.112
  • Duggan JJ, Petrie RJ (2025) The role of dynamic reciprocity in 3D cell migration: connecting cell and matrix mechanics to migratory plasticity. npj Biol Phys Mech 2:21. doi:10.1038/s44341-025-00027-1
  • Fujii Y, Ikenouchi J (2024) Cytoplasmic zoning in membrane blebs. J Biochem 175:133–140. doi:10.1093/jb/mvad084
  • Fujii Y, Sakai Y, Matsuzawa K et al. (2026) CaMKII nucleates an osmotic protein supercomplex to induce cellular bleb expansion. EMBO J 45:2433–2455. doi:10.1038/s44318-026-00703-5
  • ◆ García-Arcos JM, Jha A, Waterman CM et al. (2024a) Blebology: principles of bleb-based migration. Trends Cell Biol 34:838–853. doi:10.1016/j.tcb.2024.02.009
  • ◆ García-Arcos JM, Ziegler J, Grigolon S et al. (2024b) Rigidity percolation and active advection synergize in the actomyosin cortex to drive amoeboid cell motility. Dev Cell 59:2990–3007.e7. doi:10.1016/j.devcel.2024.06.023
  • Ghose R, Sanz-Moreno V (2026) Amoeboid cancer cells at a glance. J Cell Sci 139:jcs264674. doi:10.1242/jcs.264674
  • Goodwin K, Emrich TA, Arnold S et al. (2026) Primordial germ cells experience increasing physical confinement and DNA damage during migration in the mouse embryo. Sci Adv 12:eaec7919. doi:10.1126/sciadv.aec7919
  • Goudarzi M, Tarbashevich K, Mildner K et al. (2017) Bleb expansion in migrating cells depends on supply of membrane from cell surface invaginations. Dev Cell 43:577–587.e5. doi:10.1016/j.devcel.2017.10.030
  • Graziani V, Rodriguez-Hernandez I, Maiques O et al. (2022) The amoeboid state as part of the epithelial-to-mesenchymal transition programme. Trends Cell Biol 32:228–242. doi:10.1016/j.tcb.2021.10.004
  • Headley MB, Bins A, Nip A et al. (2016) Visualization of immediate immune responses to pioneer metastatic cells in the lung. Nature 531:513–517. doi:10.1038/nature16985
  • Ikenouchi J, Aoki K (2022) A Clockwork Bleb: cytoskeleton, calcium, and cytoplasmic fluidity. FEBS J 289:7907–7917. doi:10.1111/febs.16220
  • Jeppesen DK, Sanchez ZC, Kelley NM et al. (2025) Blebbisomes are large, organelle-rich extracellular vesicles with cell-like properties. Nat Cell Biol 27:438–448. doi:10.1038/s41556-025-01621-0
  • Jha A, Chandra A, Farahani PE et al. (2026) CD44 restricts EGFR mobility to polarize cytoskeletal signalling modules driving bleb-based migration. Nat Cell Biol 28:1408–1423. doi:10.1038/s41556-026-01981-1
  • Jia W, Czabanka M, Broggini T (2024) Cell blebbing novel therapeutic possibilities to counter metastasis. Clin Exp Metastasis 41:817–828. doi:10.1007/s10585-024-10308-z
  • Lämmermann T, Sixt M (2009) Mechanical modes of 'amoeboid' cell migration. Curr Opin Cell Biol 21:636–644. doi:10.1016/j.ceb.2009.05.003
  • Liu YJ, Le Berre M, Lautenschlaeger F et al. (2015) Confinement and low adhesion induce fast amoeboid migration of slow mesenchymal cells. Cell 160:659–672. doi:10.1016/j.cell.2015.01.007
  • Loiseau E, Schneider JAM, Keber FC et al. (2016) Shape remodeling and blebbing of active cytoskeletal vesicles. Sci Adv 2:e1500465. doi:10.1126/sciadv.1500465
  • Lopes Dos Santos R, Campillo C (2022) Studying actin-induced cell shape changes using Giant Unilamellar Vesicles and reconstituted actin networks. Biochem Soc Trans 50:1527–1539. doi:10.1042/bst20220900
  • Miyazaki M, Laboni FS, Kim T (2026) Reconstitution of actomyosin networks in cell-sized liposomes dissects distinct mechanisms of membrane blebbing and symmetry breaking. Sci Adv 12:eaed8818. doi:10.1126/sciadv.aed8818
  • Morita M, Morimoto M, Ikenouchi J et al. (2026) Bleb-based extravasation uses conserved morphodynamics but divergent calcium control. Nat Commun 17:4422. doi:10.1038/s41467-026-71052-4
  • Paluch E, Piel M, Prost J et al. (2005) Cortical actomyosin breakage triggers shape oscillations in cells and cell fragments. Biophys J 89:724–733. doi:10.1529/biophysj.105.060590
  • Petrie RJ, Yamada KM (2012) At the leading edge of three-dimensional cell migration. J Cell Sci 125:5917–5926. doi:10.1242/jcs.093732
  • Reichman-Fried M, Raz E (2023) Bleb protrusions help cancer cells to cheat death. Nature 615:402–403. doi:10.1038/d41586-023-00477-4
  • Rodriguez-Hernandez I, Maiques O, Kohlhammer L et al. (2020) WNT11-FZD7-DAAM1 signalling supports tumour initiating abilities and melanoma amoeboid invasion. Nat Commun 11:5315. doi:10.1038/s41467-020-18951-2
  • Ruprecht V, Wieser S, Callan-Jones A et al. (2015) Cortical contractility triggers a stochastic switch to fast amoeboid cell motility. Cell 160:673–685. doi:10.1016/j.cell.2015.01.008
  • Sakamoto R, Murrell MP (2024) Active tension and membrane friction mediate cortical flows and blebbing in a model actomyosin cortex. Phys Rev Res 6:033024. doi:10.1103/PhysRevResearch.6.033024
  • Schick J, Raz E (2022) Blebs—formation, regulation, positioning, and role in amoeboid cell migration. Front Cell Dev Biol 10:926394. doi:10.3389/fcell.2022.926394
  • Weems AD, Welf ES, Driscoll MK et al. (2023) Blebs promote cell survival by assembling oncogenic signalling hubs. Nature 615:517–525. doi:10.1038/s41586-023-05758-6
  • Zatulovskiy E, Tyson R, Bretschneider T et al. (2014) Bleb-driven chemotaxis of Dictyostelium cells. J Cell Biol 204:1027–1044. doi:10.1083/jcb.201306147