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Morula

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

Citations verified against PubMedNot yet clinician-reviewed10 referencesApproximately 5 min read

An early embryo that is a compact mass of about 16 blastomeres.

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 (899 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

An early embryo that is a compact mass of about 16 blastomeres. It resembles a cluster of mulberries with two types of cells, outer cells and inner cells. Morula is the stage before blastula in non-mammalian animals or a blastocyst in mammals.

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: Morulas.

Where it fits in the MeSH classification

Broader topics
Embryonic Structures

Related concepts

  • Blastomeres
  • Blastula
  • Blastocyst

Research landscape

PubMed indexes 1,348 records tagged with the MeSH heading “Morula” (counts retrieved 2026-10-31). 109 of them (8.1%) were published from 2020 onward, and the busiest period so far is through 1999 with 603 records.

Reviews account for 31 records (2.3%) and meta-analyses for 1. Randomized controlled trials number 3 (0.2%) 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: 603 records; 2000-2009: 370 records; 2010-2019: 266 records; 2020 onward: 109 records603through 19993702000-20092662010-20191092020 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 1999603
2000-2009370
2010-2019266
2020 onward109
Share of records by publication typeReviews: 2.3 percent; Meta-analyses: 0.1 percent; Randomized controlled trials: 0.2 percentReviews2.3% (31)Meta-analyses0.1% (1)Randomized controlled trials0.2% (3)
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 1,348 PubMed records
TypeRecordsShare
Reviews312.3%
Meta-analyses10.1%
Randomized controlled trials30.2%

Method: PubMed query "Morula"[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
Analysis of embryo morphokinetics, multinucleation and cleavage anomalies using continuous time-lapse monitoring in blastocyst transfer cycles
Desai N, Ploskonka S, Goodman LR, Austin C, Goldberg J, Falcone T
2014Research article
Design not recorded in structured metadata
140cc by
PMID 24951056 · DOI · PMC4074839
Tripolar mitosis and partitioning of the genome arrests human preimplantation development in vitro
Ottolini CS, Kitchen J, Xanthopoulou L, Gordon T, Summers MC, Handyside AH
2017Research article
Design not recorded in structured metadata
80cc by
PMID 28851957 · DOI · PMC5575028
Transitions in cell potency during early mouse development are driven by Notch
Menchero S, Rollan I, Lopez-Izquierdo A, Andreu MJ, Sainz de Aja J, Kang M, et al.
2019Research article
Design not recorded in structured metadata
47cc by
PMID 30958266 · DOI · PMC6486152
Clinical outcomes of co-transfer of partially and fully compacted morula versus fully compacted morula alone on day 4
Tao LL, Zheng B, Dai FF, Geng YS, Li GZ, Dai HY, et al.
2026Research article
Design not recorded in structured metadata
0cc by
PMID 42305265 · DOI · PMC13265299

Analysis of embryo morphokinetics, multinucleation and cleavage anomalies using continuous time-lapse monitoring in blastocyst transfer cycles [1]

Research article Desai N, Ploskonka S, Goodman LR, Austin C, Goldberg J, Falcone T · · 2014 · cited 140 times (Europe PMC)

Abstract reading summary — source excerpts
Opening excerpt
BackgroundTime-lapse imaging combined with embryo morphokinetics may offer a non-invasive means for improving embryo selection.
Closing excerpt
ConclusionsThese data provide us with a platform with which to potentially enhance embryo selection for transfer.
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 (341 words)

BackgroundTime-lapse imaging combined with embryo morphokinetics may offer a non-invasive means for improving embryo selection. Data from clinics worldwide are necessary to compare and ultimately develop embryo classifications models using kinetic data. The primary objective of this study was to determine if there were kinetic differences between embryos with limited potential and those more often associated with in vitro blastocyst formation and/or implantation. We also wanted to compare putative kinetic markers for embryo selection as proposed by other laboratories to what we were observing in our own laboratory setting.MethodsKinetic data and cycle outcomes were retrospectively analyzed in patients age 39 and younger with 7 or more zygotes cultured in the Embryoscope. Timing of specific events from the point of insemination were determined using time-lapse (TL) imaging. The following kinetic markers were assessed: time to syngamy (tPNf), t2, time to two cells (c), 3c (t3), 4c ( t4), 5c (t5), 8c (t8), morula (tMor), start of blastulation (tSB); tBL, blastocyst (tBL); expanded blastocyst (tEBL). Durations of the second (cc2) and third (cc3) cell cycles, the t5-t2 interval as well as time to complete synchronous divisions s1, s2 and s3 were calculated. Incidence and impact on development of nuclear and cleavage anomalies were also assessed.ResultsA total of 648 embryos transferred on day 5 were analyzed. The clinical pregnancy and implantation rate were 72% and 50%, respectively. Morphokinetic data showed that tPNf, t2,t4, t8, s1, s2,s3 and cc2 were significantly different in embryos forming blastocysts (ET or frozen) versus those with limited potential either failing to blastulate or else forming poor quality blastocysts ,ultimately discarded. Comparison of embryo kinetics in cycles with all embryos implanting (KID+) versus no implantation (KID-) suggested that markers of embryo competence to implant may be different from ability to form a blastocyst. The incidence of multinucleation and reverse cleavage amongst the embryos observed was 25% and 7%, respectively. Over 40% of embryos exhibiting these characteristics did however form blastocysts meeting our criteria for freezing.ConclusionsThese data provide us with a platform with which to potentially enhance embryo selection for transfer.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 24951056 · DOI · PMC4074839

Tripolar mitosis and partitioning of the genome arrests human preimplantation development in vitro [2]

Research article Ottolini CS, Kitchen J, Xanthopoulou L, Gordon T, Summers MC, Handyside AH · · 2017 · cited 80 times (Europe PMC)

Abstract reading summary — source excerpts
Opening excerpt
Following in vitro fertilisation (IVF), only about half of normally fertilised human embryos develop beyond cleavage and morula stages to form a blastocyst in vitro.
Closing excerpt
Failure to coordinate the cell cycle in early cleavage and regulate centrosome duplication is therefore a major cause of human preimplantation developmental arrest in vitro.
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 (195 words)

Following in vitro fertilisation (IVF), only about half of normally fertilised human embryos develop beyond cleavage and morula stages to form a blastocyst in vitro. Although many human embryos are aneuploid and genomically imbalanced, often as a result of meiotic errors inherited in the oocyte, these aneuploidies persist at the blastocyst stage and the reasons for the high incidence of developmental arrest remain unknown. Here we use genome-wide SNP genotyping and meiomapping of both polar bodies to identify maternal meiotic errors and karyomapping to fingerprint the parental chromosomes in single cells from disaggregated arrested embryos and excluded cells from blastocysts. Combined with time lapse imaging of development in culture, we demonstrate that tripolar mitoses in early cleavage cause chromosome dispersal to clones of cells with identical or closely related sub-diploid chromosome profiles resulting in intercellular partitioning of the genome. We hypothesise that following zygotic genome activation (ZGA), the combination of genomic imbalance and partial genome loss disrupts the normal pattern of embryonic gene expression blocking development at the morula-blastocyst transition. Failure to coordinate the cell cycle in early cleavage and regulate centrosome duplication is therefore a major cause of human preimplantation developmental arrest in vitro.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 28851957 · DOI · PMC5575028

Transitions in cell potency during early mouse development are driven by Notch [3]

Research article Menchero S, Rollan I, Lopez-Izquierdo A, Andreu MJ, Sainz de Aja J, Kang M, et al. · · 2019 · cited 47 times (Europe PMC)

Abstract reading summary — source excerpts
Opening excerpt
The Notch signalling pathway plays fundamental roles in diverse developmental processes in metazoans, where it is important in driving cell fate and directing differentiation of various cell types.
Closing excerpt
Our results reveal a previously undescribed role for Notch in driving transitions during the gradual loss of potency that takes place in the early mouse embryo prior to the first 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 (144 words)

The Notch signalling pathway plays fundamental roles in diverse developmental processes in metazoans, where it is important in driving cell fate and directing differentiation of various cell types. However, we still have limited knowledge about the role of Notch in early preimplantation stages of mammalian development, or how it interacts with other signalling pathways active at these stages such as Hippo. By using genetic and pharmacological tools in vivo, together with image analysis of single embryos and pluripotent cell culture, we have found that Notch is active from the 4-cell stage. Transcriptomic analysis in single morula identified novel Notch targets, such as early naïve pluripotency markers or transcriptional repressors such as TLE4. Our results reveal a previously undescribed role for Notch in driving transitions during the gradual loss of potency that takes place in the early mouse embryo prior to the first lineage decisions.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 30958266 · DOI · PMC6486152

Clinical outcomes of co-transfer of partially and fully compacted morula versus fully compacted morula alone on day 4 [4]

Research article Tao LL, Zheng B, Dai FF, Geng YS, Li GZ, Dai HY, et al. · · 2026

Abstract reading summary — source excerpts
Opening excerpt
ObjectiveTo evaluate the impact of day 4 double embryo transfer (DET) with a fully compacted morula (FCM) and a partially compacted morula (PCM) versus day 4 single embryo transfer (SET) with an FCM on clinical and neonatal outcomes in fresh cycles.
Closing excerpt
SET with FCM may be a preferable strategy for balancing clinical and multiple pregnancy rates.
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 (219 words)

ObjectiveTo evaluate the impact of day 4 double embryo transfer (DET) with a fully compacted morula (FCM) and a partially compacted morula (PCM) versus day 4 single embryo transfer (SET) with an FCM on clinical and neonatal outcomes in fresh cycles.MethodsThis was a retrospective cohort study including 889 fresh day 4 embryo transfer cycles conducted between October 2018 and December 2024. Propensity score matching (PSM) was applied to control for potential confounders and compare the clinical outcomes between SET with FCM and DET with FCM and PCM.ResultsAfter PSM, logistic regression analysis showed no significant differences in clinical pregnancy rate (CPR), miscarriage rate, live birth rate (LBR), cumulative live birth rate (CLBR), monozygotic twin rate, stillbirth rate, and cesarean section rate (CSR) (all P > 0.05). However, the DET with FCM and PCM group was associated with a significantly higher multiple pregnancy rate (MPR) (P P = 0.001) after matching. Among patients agedP > 0.05), but the MPR in the DET with FCM and PCM group was significantly higher(all PConclusionIn fresh cycles, there were no significant differences in CPR, LBR and CLBR following day 4 DET with FCM and PCM compared with SET with FCM. However, the MPR was significantly higher in the DET group. SET with FCM may be a preferable strategy for balancing clinical and multiple pregnancy rates.

Abstract reproduced from the open-access article under the recorded cc by licence; © the authors. PMID 42305265 · DOI · PMC13265299

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 The enigmatic morula: mechanisms of development, cell fate determination, self-correction and implications for ART [5]
    Coticchio G, Lagalla C, Sturmey R, Pennetta F, Borini A · Hum Reprod Update · 2019 · cited 69 times (Europe PMC)
  • Research article A review of the mechanisms and evidence for typical and atypical twinning [6]
    McNamara HC, Kane SC, Craig JM, Short RV, Umstad MP · Am J Obstet Gynecol · 2016 · cited 56 times (Europe PMC)
  • Research article Establishment of the axis in chordates: facts and speculations [7]
    Eyal-Giladi H · Development · 1997 · cited 35 times (Europe PMC)
  • Research article Regulation of present and future development by maternal regulatory signals acting on the embryo during the morula to blastocyst transition - insights from the cow [8]
    Hansen PJ, Tríbulo P · Biol Reprod · 2019 · cited 28 times (Europe PMC)
  • Research article Transcription profile in mouse four-cell, morula, and blastocyst: Genes implicated in compaction and blastocoel formation [9]
    Cui XS, Li XY, Shen XH, Bae YJ, Kang JJ, Kim NH · Mol Reprod Dev · 2007 · cited 25 times (Europe PMC)
  • Research article Membrane events associated with the generation of a blastocyst [10]
    Johnson MH · Int Rev Cytol Suppl · 1981 · cited 9 times (Europe PMC)

References

  1. Desai N, Ploskonka S, Goodman LR, Austin C, Goldberg J, Falcone T. Analysis of embryo morphokinetics, multinucleation and cleavage anomalies using continuous time-lapse monitoring in blastocyst transfer cycles. . 2014. PMID 24951056 · DOI · PMC4074839
  2. Ottolini CS, Kitchen J, Xanthopoulou L, Gordon T, Summers MC, Handyside AH. Tripolar mitosis and partitioning of the genome arrests human preimplantation development in vitro. . 2017. PMID 28851957 · DOI · PMC5575028
  3. Menchero S, Rollan I, Lopez-Izquierdo A, Andreu MJ, Sainz de Aja J, Kang M, et al.. Transitions in cell potency during early mouse development are driven by Notch. . 2019. PMID 30958266 · DOI · PMC6486152
  4. Tao LL, Zheng B, Dai FF, Geng YS, Li GZ, Dai HY, et al.. Clinical outcomes of co-transfer of partially and fully compacted morula versus fully compacted morula alone on day 4. . 2026. PMID 42305265 · DOI · PMC13265299
  5. Coticchio G, Lagalla C, Sturmey R, Pennetta F, Borini A. The enigmatic morula: mechanisms of development, cell fate determination, self-correction and implications for ART. Hum Reprod Update. 2019. PMID 30855681 · DOI
  6. McNamara HC, Kane SC, Craig JM, Short RV, Umstad MP. A review of the mechanisms and evidence for typical and atypical twinning. Am J Obstet Gynecol. 2016. PMID 26548710 · DOI
  7. Eyal-Giladi H. Establishment of the axis in chordates: facts and speculations. Development. 1997. PMID 9199355 · DOI
  8. Hansen PJ, Tríbulo P. Regulation of present and future development by maternal regulatory signals acting on the embryo during the morula to blastocyst transition - insights from the cow. Biol Reprod. 2019. PMID 31220231 · DOI · PMC8127039
  9. Cui XS, Li XY, Shen XH, Bae YJ, Kang JJ, Kim NH. Transcription profile in mouse four-cell, morula, and blastocyst: Genes implicated in compaction and blastocoel formation. Mol Reprod Dev. 2007. PMID 16998848 · DOI
  10. Johnson MH. Membrane events associated with the generation of a blastocyst. Int Rev Cytol Suppl. 1981. PMID 7019117 · DOI

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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All 542 articles in Anatomy and Physiology →

At a glance

1,348
PubMed records on this heading
8%
published since 2020
31
review articles
3
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.

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  • BlastocystAnatomy and Physiology
  • BlastomeresAnatomy and Physiology
  • GastrulaAnatomy and Physiology

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