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Gastrula

Anatomy and Physiology · NLM MeSH · updated 2026-10-31 · version 1

Citations verified against PubMedNot yet clinician-reviewed10 referencesApproximately 3 min read

The developmental stage that follows blastula or blastocyst.

Table of Contents

1. Definition2. Research landscape3. Reading the evidence4. Open-access studies5. Key reviews6. References7. About this article

Definition and research guide

This page combines a MeSH definition, publication statistics and selected research abstracts. It is a starting point for reading the literature, not a medical textbook or a systematic evidence review.

Content available: 4 complete abstracts (610 words), 10 references. Full article text is not stored or reproduced here; follow the PMID, DOI or PMC links to read the original sources and their access terms.

Definition

The developmental stage that follows blastula or blastocyst. It is characterized by the morphogenetic cell movements including invagination, ingression, and involution. Gastrulation begins with the formation of the primitive streak, and ends with the formation of three germ layers, the body plan of the mature organism.

Definition: NLM Medical Subject Headings (MeSH) scope note (Public domain (U.S. National Library of Medicine); attribution requested). View the MeSH record.

Also known as: Gastrulas; Blastopore; Blastopores; Primitive Gut; Gut, Primitive; Guts, Primitive; Primitive Guts; Gastrocoele.

Where it fits in the MeSH classification

Broader topics
Embryonic Structures

Related concepts

  • Blastula
  • Blastocyst
  • Primitive Streak
  • Germ Layers

Research landscape

PubMed indexes 3,575 records tagged with the MeSH heading “Gastrula” (counts retrieved 2026-10-31). 193 of them (5.4%) were published from 2020 onward, and the busiest period so far is through 1999 with 1,841 records.

Reviews account for 342 records (9.6%) and meta-analyses for 1. Randomized controlled trials number 0 (0.0%) as a share of all records on the topic. Randomized trials make up only a small share of this literature, so much of what is known comes from laboratory, observational or descriptive work.

PubMed records by publication periodthrough 1999: 1,841 records; 2000-2009: 1,150 records; 2010-2019: 391 records; 2020 onward: 193 records1,841through 19991,1502000-20093912010-20191932020 onward
Records in PubMed indexed with this heading, by publication period. Real counts from NCBI E-utilities; the final period covers only part of the current year. Volume reflects how much has been published, not how strong the evidence is.
Show the data
PeriodRecords
through 19991,841
2000-20091,150
2010-2019391
2020 onward193
Share of records by publication typeReviews: 9.6 percent; Meta-analyses: 0.0 percent; Randomized controlled trials: 0.0 percentReviews9.6% (342)Meta-analyses0.0% (1)Randomized controlled trials0.0% (0)
Share of all PubMed records on this heading that carry each publication type. Types overlap with other records, so the shares do not sum to 100%.
Publication-type chart data
Publication types among 3,575 PubMed records
TypeRecordsShare
Reviews3429.6%
Meta-analyses10.0%
Randomized controlled trials00.0%

Method: PubMed query "Gastrula"[MeSH Terms] with publication-type and date filters, retrieved 2026-10-31.

Reading the evidence in anatomy and physiology

Anatomy describes how body structures are built and physiology describes how they normally work, so most findings here are descriptive or mechanistic rather than evidence about treatment. Much of what is known comes from dissection, imaging, tissue studies and experiments in animals, and measurements in healthy volunteers can differ from those in people with disease or at different ages. Normal ranges vary between individuals and populations. Check whether a result was measured in people or in another species, the age and health of the participants, and the method used.

Evidence from open-access studies

These open-access studies come from PubMed’s best-match results for this heading (papers whose main topic it is), ranked here by citation count, with one recent addition. Each abstract is reproduced in full under its recorded open licence so that you can read the authors’ own words. They illustrate the literature; they are not a systematic summary of it. Citation counts measure attention, not reliability or clinical benefit.

Study comparison — selected abstracts, not a systematic review
Study / authorsYearSource label / designEurope PMC citationsLicence / source
Cortical contractility triggers a stochastic switch to fast amoeboid cell motility
Ruprecht V, Wieser S, Callan-Jones A, Smutny M, Morita H, Sako K, et al.
2015Research article
Design not recorded in structured metadata
381cc by
PMID 25679761 · DOI · PMC4328143
Single-Cell Landscape of Transcriptional Heterogeneity and Cell Fate Decisions during Mouse Early Gastrulation
Mohammed H, Hernando-Herraez I, Savino A, Scialdone A, Macaulay I, Mulas C, et al.
2017Research article
Design not recorded in structured metadata
334cc by
PMID 28768204 · DOI · PMC5554778
Anteroposterior polarity and elongation in the absence of extra-embryonic tissues and of spatially localised signalling in gastruloids: mammalian embryonic organoids
Turner DA, Girgin M, Alonso-Crisostomo L, Trivedi V, Baillie-Johnson P, Glodowski CR, et al.
2017Research article
Design not recorded in structured metadata
258cc by
PMID 28951435 · DOI · PMC5702072
Notch coordinates self-organization of germ layers and axial polarity in sea anemone gastruloids
Narayanaswamy S, Haas F, Haillot E, Cole AG, Tanaka EM, Technau U
2026Research article
Design not recorded in structured metadata
1cc by
PMID 42321198 · DOI · PMC13370024

Cortical contractility triggers a stochastic switch to fast amoeboid cell motility [1]

Research article Ruprecht V, Wieser S, Callan-Jones A, Smutny M, Morita H, Sako K, et al. · · 2015 · cited 381 times (Europe PMC)

Abstract reading summary — source excerpts
Opening excerpt
3D amoeboid cell migration is central to many developmental and disease-related processes such as cancer metastasis.
Closing excerpt
We further show that rearward cortical flows drive stable-bleb cell migration in various adhesive and non-adhesive environments, unraveling a highly versatile amoeboid migration phenotype.
Interpretation limits
Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (136 words)

3D amoeboid cell migration is central to many developmental and disease-related processes such as cancer metastasis. Here, we identify a unique prototypic amoeboid cell migration mode in early zebrafish embryos, termed stable-bleb migration. Stable-bleb cells display an invariant polarized balloon-like shape with exceptional migration speed and persistence. Progenitor cells can be reversibly transformed into stable-bleb cells irrespective of their primary fate and motile characteristics by increasing myosin II activity through biochemical or mechanical stimuli. Using a combination of theory and experiments, we show that, in stable-bleb cells, cortical contractility fluctuations trigger a stochastic switch into amoeboid motility, and a positive feedback between cortical flows and gradients in contractility maintains stable-bleb cell polarization. We further show that rearward cortical flows drive stable-bleb cell migration in various adhesive and non-adhesive environments, unraveling a highly versatile amoeboid migration phenotype.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 25679761 · DOI · PMC4328143

Single-Cell Landscape of Transcriptional Heterogeneity and Cell Fate Decisions during Mouse Early Gastrulation [2]

Research article Mohammed H, Hernando-Herraez I, Savino A, Scialdone A, Macaulay I, Mulas C, et al. · · 2017 · cited 334 times (Europe PMC)

Abstract reading summary — source excerpts
Opening excerpt
The mouse inner cell mass (ICM) segregates into the epiblast and primitive endoderm (PrE) lineages coincident with implantation of the embryo.
Closing excerpt
Our study systematically charts transcriptional noise and uncovers molecular processes associated with early lineage decisions.
Interpretation limits
Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (153 words)

The mouse inner cell mass (ICM) segregates into the epiblast and primitive endoderm (PrE) lineages coincident with implantation of the embryo. The epiblast subsequently undergoes considerable expansion of cell numbers prior to gastrulation. To investigate underlying regulatory principles, we performed systematic single-cell RNA sequencing (seq) of conceptuses from E3.5 to E6.5. The epiblast shows reactivation and subsequent inactivation of the X chromosome, with Zfp57 expression associated with reactivation and inactivation together with other candidate regulators. At E6.5, the transition from epiblast to primitive streak is linked with decreased expression of polycomb subunits, suggesting a key regulatory role. Notably, our analyses suggest elevated transcriptional noise at E3.5 and within the non-committed epiblast at E6.5, coinciding with exit from pluripotency. By contrast, E6.5 primitive streak cells became highly synchronized and exhibit a shortened G1 cell-cycle phase, consistent with accelerated proliferation. Our study systematically charts transcriptional noise and uncovers molecular processes associated with early lineage decisions.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 28768204 · DOI · PMC5554778

Anteroposterior polarity and elongation in the absence of extra-embryonic tissues and of spatially localised signalling in gastruloids: mammalian embryonic organoids [3]

Research article Turner DA, Girgin M, Alonso-Crisostomo L, Trivedi V, Baillie-Johnson P, Glodowski CR, et al. · · 2017 · cited 258 times (Europe PMC)

Abstract reading summary — source excerpts
Opening excerpt
The establishment of the anteroposterior (AP) axis is a crucial step during animal embryo development.
Closing excerpt
Furthermore, we suggest that Wnt signalling has a separable activity involved in the elongation of the axis.
Interpretation limits
Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (181 words)

The establishment of the anteroposterior (AP) axis is a crucial step during animal embryo development. In mammals, genetic studies have shown that this process relies on signals spatiotemporally deployed in the extra-embryonic tissues that locate the position of the head and the onset of gastrulation, marked by T/Brachyury (T/Bra) at the posterior of the embryo. Here, we use gastruloids, mESC-based organoids, as a model system with which to study this process. We find that gastruloids localise T/Bra expression to one end and undergo elongation similar to the posterior region of the embryo, suggesting that they develop an AP axis. This process relies on precisely timed interactions between Wnt/β-catenin and Nodal signalling, whereas BMP signalling is dispensable. Additionally, polarised T/Bra expression occurs in the absence of extra-embryonic tissues or localised sources of signals. We suggest that the role of extra-embryonic tissues in the mammalian embryo might not be to induce the axes but to bias an intrinsic ability of the embryo to initially break symmetry. Furthermore, we suggest that Wnt signalling has a separable activity involved in the elongation of the axis.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 28951435 · DOI · PMC5702072

Notch coordinates self-organization of germ layers and axial polarity in sea anemone gastruloids [4]

Research article Narayanaswamy S, Haas F, Haillot E, Cole AG, Tanaka EM, Technau U · · 2026 · cited 1 times (Europe PMC)

Abstract reading summary — source excerpts
Opening excerpt
Reaggregated early gastrula cells ("gastruloids") of the sea anemone Nematostella vectensis are able to regenerate into whole polyps within a few days.
Closing excerpt
This demonstrates how adoption of ancestral regulatory networks enables a morphospace converging to similar body plans, thus contributing to evolutionary robustness.
Interpretation limits
Verbatim excerpts from the authors’ abstract; the closing excerpt may describe a finding, limitation or recommendation. Design, population, effect size and uncertainty have not been independently extracted or appraised. Read the complete abstract and linked paper before interpreting the findings.
Read complete abstract (140 words)

Reaggregated early gastrula cells ("gastruloids") of the sea anemone Nematostella vectensis are able to regenerate into whole polyps within a few days. However, the cellular and molecular mechanisms underlying restoration of body axis and germ layers remains largely unknown. Here, we show that mesodermal cells sort to the periphery of the gastruloid, where they form cell clusters with hitchhiking endodermal cells. One of these clusters immigrates at one pole, forming the future pharynx and inner layer. This morphogenetic behavior that enables the symmetry break of the organizer tissue, requires a Wnt-Notch signaling feedback loop. This highlights a hitherto unknown role of Notch signaling in self-organizing gastruloids. Conservation of Notch-mediated boundary formation between germ layers mirrors similar mechanisms in bilaterians. This demonstrates how adoption of ancestral regulatory networks enables a morphospace converging to similar body plans, thus contributing to evolutionary robustness.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 42321198 · DOI · PMC13370024

Key reviews and meta-analyses

The most cited review-type papers among PubMed’s best-match results for this heading, listed with bibliographic details only. Follow the links for the full text and access terms.

  • Research article Gene function in mouse embryogenesis: get set for gastrulation [5]
    Tam PP, Loebel DA · Nat Rev Genet · 2007 · cited 470 times (Europe PMC)
  • Research article Mouse gastrulation: the formation of a mammalian body plan [6]
    Tam PP, Behringer RR · Mech Dev · 1997 · cited 465 times (Europe PMC)
  • Research article How we are shaped: the biomechanics of gastrulation [7]
    Keller R, Davidson LA, Shook DR · Differentiation · 2003 · cited 344 times (Europe PMC)
  • Research article Conserved patterns of cell movements during vertebrate gastrulation [8]
    Solnica-Krezel L · Curr Biol · 2005 · cited 300 times (Europe PMC)
  • Research article Planar cell polarization: do the same mechanisms regulate Drosophila tissue polarity and vertebrate gastrulation? [9]
    Mlodzik M · Trends Genet · 2002 · cited 238 times (Europe PMC)
  • Research article From egg to gastrula: how the cell cycle is remodeled during the Drosophila mid-blastula transition [10]
    Farrell JA, O'Farrell PH · Annu Rev Genet · 2014 · cited 182 times (Europe PMC)

References

  1. Ruprecht V, Wieser S, Callan-Jones A, Smutny M, Morita H, Sako K, et al.. Cortical contractility triggers a stochastic switch to fast amoeboid cell motility. . 2015. PMID 25679761 · DOI · PMC4328143
  2. Mohammed H, Hernando-Herraez I, Savino A, Scialdone A, Macaulay I, Mulas C, et al.. Single-Cell Landscape of Transcriptional Heterogeneity and Cell Fate Decisions during Mouse Early Gastrulation. . 2017. PMID 28768204 · DOI · PMC5554778
  3. Turner DA, Girgin M, Alonso-Crisostomo L, Trivedi V, Baillie-Johnson P, Glodowski CR, et al.. Anteroposterior polarity and elongation in the absence of extra-embryonic tissues and of spatially localised signalling in gastruloids: mammalian embryonic organoids. . 2017. PMID 28951435 · DOI · PMC5702072
  4. Narayanaswamy S, Haas F, Haillot E, Cole AG, Tanaka EM, Technau U. Notch coordinates self-organization of germ layers and axial polarity in sea anemone gastruloids. . 2026. PMID 42321198 · DOI · PMC13370024
  5. Tam PP, Loebel DA. Gene function in mouse embryogenesis: get set for gastrulation. Nat Rev Genet. 2007. PMID 17387317 · DOI
  6. Tam PP, Behringer RR. Mouse gastrulation: the formation of a mammalian body plan. Mech Dev. 1997. PMID 9431800 · DOI
  7. Keller R, Davidson LA, Shook DR. How we are shaped: the biomechanics of gastrulation. Differentiation. 2003. PMID 12694202 · DOI
  8. Solnica-Krezel L. Conserved patterns of cell movements during vertebrate gastrulation. Curr Biol. 2005. PMID 15797016 · DOI
  9. Mlodzik M. Planar cell polarization: do the same mechanisms regulate Drosophila tissue polarity and vertebrate gastrulation?. Trends Genet. 2002. PMID 12414186 · DOI
  10. Farrell JA, O'Farrell PH. From egg to gastrula: how the cell cycle is remodeled during the Drosophila mid-blastula transition. Annu Rev Genet. 2014. PMID 25195504 · DOI · PMC4484755

About this article

Sources
NLM Medical Subject Headings (definition, synonyms); NCBI PubMed (counts, identifiers); Europe PMC (open-access abstracts and citation counts).
Verification
Every reference was resolved against PubMed on 2026-10-31; title, year and DOI matched.
Review state
automated source verified; no human expert review has been recorded yet.
Version
1, last updated 2026-10-31.

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Related research topics

  • GastrulationAnatomy and Physiology
  • MorulaAnatomy and Physiology
  • Neural CrestAnatomy and Physiology
  • BlastulaAnatomy and Physiology
  • BlastocystAnatomy and Physiology
  • TrophoblastsAnatomy and Physiology

All 542 articles in Anatomy and Physiology →

At a glance

3,575
PubMed records on this heading
5%
published since 2020
342
review articles
0
randomized trials
10
verified references below

Counts come from PubMed on 2026-10-31; they show how much has been written, not how strong the evidence is.

Related topics See all →

  • GastrulationAnatomy and Physiology
  • MorulaAnatomy and Physiology
  • Neural CrestAnatomy and Physiology
  • BlastulaAnatomy and Physiology
  • BlastocystAnatomy and Physiology

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