Morula Development

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Introduction

Human morula (day 2)[1]
Human morula (day 3)[1]

(Latin, morula = mulberry) An early stage in post-fertilization development when cells have rapidly mitotically divided to produce a solid mass of cells (12-15 cells) with a "mulberry" appearance. This stage is followed by formation of a cavity in this cellular mass blastocyst stage.

A key event prior to morula formation is "compaction", where the 8 cell embryo undergoes changes in cell morphology and cell-cell adhesion that initiates the formation of this solid ball of cells.


In humans, morula stage of development occurs during the first days in week one following fertilization (GA week 3) and is described as Carnegie stage 2. This stage is followed by formation of a cavity, the blastocoel, which defines formation of the blastocyst.

ART Preimplantation blastomere biopsy
ART Preimplantation blastomere biopsy[2]


In Assisted Reproductive Technology, the morula stage is when one of the earliest prenatal diagnostic test can be carried out, by removing a single cell (blastomere) and carrying out genetic diagnosis on its DNA.
Morula Links: morula | Carnegie stage 2 | mitosis | blastocyst | fertilization | Week 1 | Lecture - Week 1 | Category:Carnegie Stage 2 | Category:Morula

Some Recent Findings

  • Diversity of human and mouse homeobox gene expression in development and adult tissues[3] "8-cell to morula, 12 eutherian-specific homeobox genes not detectable outside of reproductive tissues or the embryo (RHOXF2, RHOXF2B, CPHX1, CPHX2, DPRX, LEUTX, TPRX1, TPRX2, ARGFX, NANOGNB, DUXA, DUXB). Hox
  • Par-aPKC-dependent and -independent mechanisms cooperatively control cell polarity, Hippo signaling, and cell positioning in 16-cell stage mouse embryos[4] "In preimplantation mouse embryos, the Hippo signaling pathway plays a central role in regulating the fates of the trophectoderm (TE) and the inner cell mass (ICM). In early blastocysts with more than 32 cells, the Par-aPKC system controls polarization of the outer cells along the apicobasal axis, and cell polarity suppresses Hippo signaling." Hippo | Mouse Development
  • Functional genomics of 5- to 8-cell stage human embryos by blastomere single-cell cDNA analysis[5] "Forty-nine blastomeres from 5- to 8-cell human embryos have been investigated following an efficient single-cell cDNA amplification protocol to provide a template for high-density microarray analysis. The previously described markers, characteristic of Inner Cell Mass (ICM) (n = 120), stemness (n = 190) and Trophectoderm (TE) (n = 45), were analyzed, and a housekeeping pattern of 46 genes was established. ...In summary, the global single-cell cDNA amplification microarray analysis of the 5- to 8-cell stage human embryos reveals that blastomere fate is not committed to ICM or TE."
  • Non-invasive imaging of human embryos before embryonic genome activation predicts development to the blastocyst stage[6] "We report studies of preimplantation human embryo development that correlate time-lapse image analysis and gene expression profiling. By examining a large set of zygotes from in vitro fertilization (IVF), we find that success in progression to the blastocyst stage can be predicted with >93% sensitivity and specificity by measuring three dynamic, noninvasive imaging parameters by day 2 after fertilization, before embryonic genome activation (EGA)."
More recent papers  
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This table shows an automated computer PubMed search using the listed sub-heading term.

  • Therefore the list of references do not reflect any editorial selection of material based on content or relevance.
  • References appear in this list based upon the date of the actual page viewing.

References listed on the rest of the content page and the associated discussion page (listed under the publication year sub-headings) do include some editorial selection based upon both relevance and availability.

Links: References | Discussion Page | Pubmed Most Recent | Journal Searches


Search term: Morula Development

Jigal Haas, Jim Meriano, Rawad Bassil, Eran Barzilay, Eran Zilberberg, Robert F Casper Developmental potential of slow-developing embryos: day-5 morulae compared with day-5 cavitating morulae. Fertil. Steril.: 2018; PubMed 30442415

Jing Xu, Maralee S Lawson, Shoukhrat M Mitalipov, Byung S Park, Fuhua Xu Stage-specific modulation of antimüllerian hormone promotes primate follicular development and oocyte maturation in the matrix-free three-dimensional culture. Fertil. Steril.: 2018, 110(6);1162-1172 PubMed 30396561

Daniel Bodri, Robert Milewski, Jazmina Yao Serna, Takeshi Sugimoto, Ryutaro Kato, Tsunekazu Matsumoto, Satoshi Kawachiya Predicting live birth by combining cleavage and blastocyst-stage time-lapse variables using a hierarchical and a data mining-based statistical model. Reprod Biol: 2018; PubMed 30389297

W T M Soliman, K Gh M Mahmoud, A R M El-Khawagah, M M M Kandiel, M E A Abouel-Roos, A E Abdel-Ghaffar, A E I El Azab ##Title## Iran J Vet Res: 2018, 19(3);178-181 PubMed 30349563

S Miranda, N Carolino, H Vilhena, R Payan-Carreira, R M L N Pereira Early embryo development, number, quality, and location and the relationship with plasma progesterone in dogs. Anim. Reprod. Sci.: 2018; PubMed 30322674

Movies

Model embryo to 32 cell stage icon.jpg
 ‎‎Morula Model
Page | Play
Mouse zygote division icon.jpg
 ‎‎Zygote Mitosis
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Mouse zygote division 02 icon.jpg
 ‎‎Early Division
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Parental genome mix 01 icon.jpg
 ‎‎Parental Genomes
Page | Play
Mouse blastocyst movie icon.jpg
 ‎‎Mouse Blastocyst
Page | Play

Movies

Compaction

  • E-cadherin mediated adhesion initiates at compaction at the 8-cell stage
  • regulated post-translationally via protein kinase C and other signalling molecules

Blastomere Division

Spindle orientation calculation.[7]

An in vitro study of human blastocyst development[8] showed that those blastomeres that initially divide quickly are more likely to develop to blastocyst stage.

A recent study in mice showed that there was no specific orientation of the mitotic spindle during cell division in the 8 to 16 cell stage transition.[7] This suggests no predetermined cleavage pattern (pre-patterned) at the 8 cell stage and only modulated by the extent of cell rounding up during mitosis. In other species, such as the worm C.elegans and ascidians, have specific patterns of spindle orientation from the zygote stage.

Model Human Morula Development

The following figure is from a recent study[6] using video and genetic analysis of in vitro human development during week 1 following fertilization.

Model human blastocyst development.jpg

  • EGA - embryonic genome activation
  • ESSP - embryonic stage–specific pattern, four unique embryonic stage–specific patterns (1-4)
Links: Figure with legend


Morulas in Other Species

Mouse Morula


Links: Mouse Development

Sea Urchin Morula

Sea Urchin- early embryo cleavage pattern.jpg

Sea Urchin early embryo cleavage pattern (SDB Gallery Images)


Links: Sea Urchin Development

Bovine Morula

Bovine morula 01.jpg

Bovine Morula[9]

  • Image shows DNA staining (white) and f-actin filaments (orange) at day 4. Scale bars represent 100 µm.
  • Pale staining round nuclei are at interphase.
  • Arrow shows single nucleus at prophase.
  • A single nucleus is seen at metaphase.
  • Condensed bright nuclei are apoptotic.


Links: Bovine Development | Mitosis

Morula Biopsy

Morula biopsy 01.jpg

Biopsy of compact morula-stage embryos[10]

  • (A) A compact morula-stage embryo before biopsy.
  • (B–G) Steps of the biopsy.
  • (H) An embryo at 2 h after biopsy.


Links: Prenatal Diagnosis | Assisted Reproductive Technology

References

  1. 1.0 1.1 Zhang P, Zucchelli M, Bruce S, Hambiliki F, Stavreus-Evers A, Levkov L, Skottman H, Kerkelä E, Kere J & Hovatta O. (2009). Transcriptome profiling of human pre-implantation development. PLoS ONE , 4, e7844. PMID: 19924284 DOI.
  2. Milachich T. (2014). New advances of preimplantation and prenatal genetic screening and noninvasive testing as a potential predictor of health status of babies. Biomed Res Int , 2014, 306505. PMID: 24783200 DOI.
  3. Dunwell TL & Holland PW. (2016). Diversity of human and mouse homeobox gene expression in development and adult tissues. BMC Dev. Biol. , 16, 40. PMID: 27809766 DOI.
  4. Hirate Y, Hirahara S, Inoue K, Kiyonari H, Niwa H & Sasaki H. (2015). Par-aPKC-dependent and -independent mechanisms cooperatively control cell polarity, Hippo signaling, and cell positioning in 16-cell stage mouse embryos. Dev. Growth Differ. , 57, 544-56. PMID: 26450797 DOI.
  5. Galán A, Montaner D, Póo ME, Valbuena D, Ruiz V, Aguilar C, Dopazo J & Simón C. (2010). Functional genomics of 5- to 8-cell stage human embryos by blastomere single-cell cDNA analysis. PLoS ONE , 5, e13615. PMID: 21049019 DOI.
  6. 6.0 6.1 Wong CC, Loewke KE, Bossert NL, Behr B, De Jonge CJ, Baer TM & Reijo Pera RA. (2010). Non-invasive imaging of human embryos before embryonic genome activation predicts development to the blastocyst stage. Nat. Biotechnol. , 28, 1115-21. PMID: 20890283 DOI.
  7. 7.0 7.1 Dard N, Louvet-Vallée S & Maro B. (2009). Orientation of mitotic spindles during the 8- to 16-cell stage transition in mouse embryos. PLoS ONE , 4, e8171. PMID: 19997595 DOI.
  8. Fenwick J, Platteau P, Murdoch AP & Herbert M. (2002). Time from insemination to first cleavage predicts developmental competence of human preimplantation embryos in vitro. Hum. Reprod. , 17, 407-12. PMID: 11821286
  9. Leidenfrost S, Boelhauve M, Reichenbach M, Güngör T, Reichenbach HD, Sinowatz F, Wolf E & Habermann FA. (2011). Cell arrest and cell death in mammalian preimplantation development: lessons from the bovine model. PLoS ONE , 6, e22121. PMID: 21811561 DOI.
  10. Zakharova EE, Zaletova VV & Krivokharchenko AS. (2014). Biopsy of human morula-stage embryos: outcome of 215 IVF/ICSI cycles with PGS. PLoS ONE , 9, e106433. PMID: 25191937 DOI.

Reviewss

Articles

Bessonnard S, Mesnard D & Constam DB. (2015). PC7 and the related proteases Furin and Pace4 regulate E-cadherin function during blastocyst formation. J. Cell Biol. , 210, 1185-97. PMID: 26416966 DOI.

Dzamba BJ, Jakab KR, Marsden M, Schwartz MA & DeSimone DW. (2009). Cadherin adhesion, tissue tension, and noncanonical Wnt signaling regulate fibronectin matrix organization. Dev. Cell , 16, 421-32. PMID: 19289087 DOI.

Santos J, Pereira CF, Di-Gregorio A, Spruce T, Alder O, Rodriguez T, Azuara V, Merkenschlager M & Fisher AG. (2010). Differences in the epigenetic and reprogramming properties of pluripotent and extra-embryonic stem cells implicate chromatin remodelling as an important early event in the developing mouse embryo. Epigenetics Chromatin , 3, 1. PMID: 20157423 DOI.

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Cite this page: Hill, M.A. (2018, November 19) Embryology Morula Development. Retrieved from https://embryology.med.unsw.edu.au/embryology/index.php/Morula_Development

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© Dr Mark Hill 2018, UNSW Embryology ISBN: 978 0 7334 2609 4 - UNSW CRICOS Provider Code No. 00098G