|Embryology - 19 Jan 2018 Expand to Translate|
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(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 the first week following fertilization and is described as Carnegie stage 2. This stage is followed by formation of a cavity, the blastocoel, which defines formation of the blastocyst.
||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
|More recent papers|
This table shows an automated computer PubMed search using the listed sub-heading term.
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.
Hakimeh Akbari, Seyed Hassan Eftekhar Vaghefi, Abbas Shahedi, Victoria Habibzadeh, Tooraj Reza Mirshekari, Aboozar Ganjizadegan, Hamidreza Mollaei, Meysam Ahmadi, Seyed Noureddin Nematollahi-Mahani Mesenchymal Stem Cell-Conditioned Medium Modulates Apoptotic and Stress-Related Gene Expression, Ameliorates Maturation and Allows for the Development of Immature Human Oocytes after Artificial Activation. Genes (Basel): 2017, 8(12); PubMed 29292728
Yawei Gao, Xiaoyu Liu, Bin Tang, Chong Li, Zhaohui Kou, Lin Li, Wenqiang Liu, You Wu, Xiaochen Kou, Jingyi Li, Yanhong Zhao, Jiqing Yin, Hong Wang, She Chen, Lujian Liao, Shaorong Gao Protein Expression Landscape of Mouse Embryos during Pre-implantation Development. Cell Rep: 2017, 21(13);3957-3969 PubMed 29281840
Maryam Zand-Vakili, Afsaneh Golkar-Narenji, Paul E Mozdziak, Hussein Eimani An in vitro study on oocyte and follicles of transplanted ovaries treated with vascular endothelial growth factor. J Turk Ger Gynecol Assoc: 2017, 18(4);167-173 PubMed 29278228
Julia M Fraile, Diana Campos-Iglesias, Francisco Rodríguez, Aurora Astudillo, Roser Vilarrasa-Blasi, Nuria Verdaguer-Dot, Miguel A Prado, Joao A Paulo, Steven P Gygi, José I Martín-Subero, José M P Freije, Carlos López-Otín Loss of the Deubiquitinase USP36 Destabilizes the RNA Helicase DHX33 and Causes Preimplantation Lethality in Mice. J. Biol. Chem.: 2017; PubMed 29273634
Verónica M Negrón-Pérez, Peter J Hansen Role of Yes-Associated Protein 1, Angiomotin and Mitogen Activated Kinase Kinase 1/2 in Development of the Bovine Blastocyst. Biol. Reprod.: 2017; PubMed 29228123
- E-cadherin mediated adhesion initiates at compaction at the 8-cell stage
- regulated post-translationally via protein kinase C and other signalling molecules
An in vitro study of human blastocyst development 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. 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 using video and genetic analysis of in vitro human development during week 1 following fertilization.
- 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
- Links: Mouse Development
Sea Urchin Morula
Sea Urchin early embryo cleavage pattern (SDB Gallery Images)
- Links: Sea Urchin Development
- 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.
- Pu Zhang, Marco Zucchelli, Sara Bruce, Fredwell Hambiliki, Anneli Stavreus-Evers, Lev Levkov, Heli Skottman, Erja Kerkelä, Juha Kere, Outi Hovatta Transcriptome profiling of human pre-implantation development. PLoS ONE: 2009, 4(11);e7844 PubMed 19924284 | PMC2773928 | PLoS One
- Tanya Milachich New advances of preimplantation and prenatal genetic screening and noninvasive testing as a potential predictor of health status of babies. Biomed Res Int: 2014, 2014;306505 PubMed 24783200 | PMC3982254 | Biomed Res Int.
- Thomas L Dunwell, Peter W H Holland Diversity of human and mouse homeobox gene expression in development and adult tissues. BMC Dev. Biol.: 2016, 16(1);40 PubMed 27809766
- Yoshikazu Hirate, Shino Hirahara, Ken-Ichi Inoue, Hiroshi Kiyonari, Hiroshi Niwa, Hiroshi Sasaki Par-aPKC-dependent and -independent mechanisms cooperatively control cell polarity, Hippo signaling, and cell positioning in 16-cell stage mouse embryos. Dev. Growth Differ.: 2015; PubMed 26450797
- Galán A, Montaner D, Póo ME, Valbuena D, Ruiz V, Aguilar C, Dopazo J, Simón C. Functional genomics of 5- to 8-cell stage human embryos by blastomere single-cell cDNA analysis. PLoS One. 2010 Oct 26;5(10):e13615. PMID21049019 | PLoS One.
- Connie C Wong, Kevin E Loewke, Nancy L Bossert, Barry Behr, Christopher J De Jonge, Thomas M Baer, Renee A Reijo Pera Non-invasive imaging of human embryos before embryonic genome activation predicts development to the blastocyst stage. Nat. Biotechnol.: 2010, 28(10);1115-21 PubMed 20890283 | Nat Biotechnol.
- Nicolas Dard, Sophie Louvet-Vallée, Bernard Maro Orientation of mitotic spindles during the 8- to 16-cell stage transition in mouse embryos. PLoS ONE: 2009, 4(12);e8171 PubMed 19997595 | PLoS One.
- J Fenwick, P Platteau, A P Murdoch, M Herbert Time from insemination to first cleavage predicts developmental competence of human preimplantation embryos in vitro. Hum. Reprod.: 2002, 17(2);407-12 PubMed 11821286
- Sandra Leidenfrost, Marc Boelhauve, Myriam Reichenbach, Tuna Güngör, Horst-Dieter Reichenbach, Fred Sinowatz, Eckhard Wolf, Felix A Habermann Cell arrest and cell death in mammalian preimplantation development: lessons from the bovine model. PLoS ONE: 2011, 6(7);e22121 PubMed 21811561 | PLoS One.
Sylvain Bessonnard, Daniel Mesnard, Daniel B Constam PC7 and the related proteases Furin and Pace4 regulate E-cadherin function during blastocyst formation. J. Cell Biol.: 2015, 210(7);1185-97 PubMed 26416966
Bette J Dzamba, Karoly R Jakab, Mungo Marsden, Martin A Schwartz, Douglas W DeSimone Cadherin adhesion, tissue tension, and noncanonical Wnt signaling regulate fibronectin matrix organization. Dev. Cell: 2009, 16(3);421-32 PubMed 19289087
Joana Santos, C Filipe Pereira, Aida Di-Gregorio, Thomas Spruce, Olivia Alder, Tristan Rodriguez, Véronique Azuara, Matthias Merkenschlager, Amanda G Fisher 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: 2010, 3;1 PubMed 20157423
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Cite this page: Hill, M.A. 2018 Embryology Morula. Retrieved January 19, 2018, from https://embryology.med.unsw.edu.au/embryology/index.php/Morula
- © Dr Mark Hill 2018, UNSW Embryology ISBN: 978 0 7334 2609 4 - UNSW CRICOS Provider Code No. 00098G