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| {{Header}}
| | # REDIRECT [[Morula Development]] |
| ==Introduction==
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| [[File:Human embryo day 2.jpg|thumb|300px|Human morula (day 2){{#pmid:19924284|PMID19924284}}]]
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| [[File:Human_embryo_day_3.jpg|thumb|300px|Human morula (day 3){{#pmid:19924284|PMID19924284}}]]
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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.
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| 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.
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| 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]].
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| {|
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| | [[File:Preimplantation blastomere biopsy.jpg|thumb|150px|alt=ART Preimplantation blastomere biopsy|ART Preimplantation blastomere biopsy{{#pmid:24783200|PMID24783200}}]]
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| | 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.
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| {{Morula Links}}
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| == Some Recent Findings ==
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| {|
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| |-bgcolor="F5FAFF"
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| * D'''iversity of human and mouse homeobox gene expression in development and adult tissues'''{{#pmid:27809766|PMID27809766}} "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). [[Developmental Signals - Homeobox|Hox]]
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| * '''Par-aPKC-dependent and -independent mechanisms cooperatively control cell polarity, Hippo signaling, and cell positioning in 16-cell stage mouse embryos'''{{#pmid:26450797|PMID26450797}} "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." [[Developmental Signals - Hippo|Hippo]] | [[Mouse Development]]
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| * '''Functional genomics of 5- to 8-cell stage human embryos by blastomere single-cell cDNA analysis'''{{#pmid:21049019|PMID21049019}} "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."
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| * '''Non-invasive imaging of human embryos before embryonic genome activation predicts development to the blastocyst stage'''{{#pmid:20890283|PMID20890283}} "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)."
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| |}
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| {| class="wikitable mw-collapsible mw-collapsed"
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| ! More recent papers
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| | [[File:Mark_Hill.jpg|90px|left]] {{Most_Recent_Refs}}
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| Search term: ''Morula Development''
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| <pubmed limit=5>Morula Development</pubmed>
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| |}
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| ==Movies==
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| {|
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| | valign="bottom"|{{Morula model movie}}
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| | valign="bottom"|{{Mouse zygote movie}}
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| | valign="bottom"|{{Mouse zygote movie 1}}
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| | valign="bottom"|{{Mouse zygote movie 2}}
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| | valign="bottom"|{{Mouse Blastocyst movie}}
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| |}
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| [[Movies]]
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| ==Compaction==
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| * E-cadherin mediated adhesion initiates at compaction at the 8-cell stage
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| * regulated post-translationally via protein kinase C and other signalling molecules
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| ==Blastomere Division==
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| [[File:Blastomere mitotic spindle orientation.jpg|thumb|Spindle orientation calculation.{{#pmid:19997595|PMID19997595}}]]
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| An in vitro study of human blastocyst development{{#pmid:11821286|PMID11821286}} showed that those blastomeres that initially divide quickly are more likely to develop to blastocyst stage.
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| 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.{{#pmid:19997595|PMID19997595}} 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.
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| ==Model Human Morula Development==
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| The following figure is from a recent study{{#pmid:20890283|PMID20890283}} using video and genetic analysis of ''in vitro'' human development during week 1 following fertilization.
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| [[File:Model human blastocyst development.jpg|800px]]
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| * EGA - embryonic genome activation
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| * ESSP - embryonic stage–specific pattern, four unique embryonic stage–specific patterns (1-4)
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| :'''Links:''' [[:File:Model human blastocyst development.jpg|Figure with legend]]
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| ==Morulas in Other Species==
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| ===Mouse Morula===
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| <gallery>
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| File:Mouse-morula_01.jpg|4 cell morula stage development
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| File:Mouse_Sox2_expression_01.jpg|Sox2 expression
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| File:Mouse- preimplantation gene expression 02.jpg|Early gene expression
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| File:Mouse- preimplantation gene expression 03.jpg|Early gene expression
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| File:Mouse- preimplantation gene expression 04.jpg|Early gene expression
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| File:Mouse- preimplantation gene expression 05.jpg|Early gene expression
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| </gallery>
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| :'''Links:''' [[Mouse Development]]
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| ===Sea Urchin Morula===
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| [[File:Sea Urchin- early embryo cleavage pattern.jpg|300px]]
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| '''Sea Urchin''' early embryo cleavage pattern (SDB Gallery Images)
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| :'''Links:''' [[Sea Urchin Development]]
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| ===Bovine Morula===
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| [[File:Bovine_morula_01.jpg|600px]]
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| '''Bovine Morula'''{{#pmid:21811561|PMID21811561}}
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| * Image shows DNA staining (white) and f-actin filaments (orange) at day 4. Scale bars represent 100 µm.
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| * Pale staining round nuclei are at interphase.
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| * Arrow shows single nucleus at prophase.
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| * A single nucleus is seen at metaphase.
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| * Condensed bright nuclei are apoptotic.
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| :'''Links:''' [[Bovine Development]] | [[Cell Division - Mitosis|Mitosis]]
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| ==References==
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| <references/>
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| ===Reviewss===
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| ===Articles===
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| {{#pmid:26416966}}
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| {{#pmid:19289087}}
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| {{#pmid:20157423}}
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| ===Search PubMed===
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| '''Search Pubmed:''' [http://www.ncbi.nlm.nih.gov/sites/entrez?db=pubmed&cmd=search&term=morula%20development morula development] | [http://www.ncbi.nlm.nih.gov/sites/entrez?db=pubmed&cmd=search&term=blastomere%20development blastomere development] |
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| {{Glossary}}
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| {{Footer}}
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| [[Category:Human Embryo]] [[Category:Carnegie Stage 2]] [[Category:Week 1]] [[Category:Morula]]
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