These notes cover the fourth week of embryonic development, which is the beginning of organogenesis, (specific tissues and systems are beginning to differentiate) from the trilaminar embryo. On the embryo surface sensory placodes and limb buds appear. Sensory placodes (otic, lens, nasal) will form specific components of the ear, eye and nose. Limb buds form from ectoderm and mesoderm (somite) components and are the "paddle-like" projections from the trunk which will form all the upper and lower limb components. Within the embryo, this period of organogensis is usually extended to cover until 8 weeks of development. Folding of the embryo continues and the earliest functioning organ is the heart. Other systems such as the circulatory, digestive, urogenital and nervous system begin to all take shape. As systems are beginning to develop, each page in this section is only a brief summary with additional links to specific notes covering these systems/tissues. Note that most tissue development during week 4 are covered in System Development notes. |
Embryo Development Week 4 |
Page Links: Introduction | Some Recent Findings | Reading | Objectives | Computer Activities | Development Overview | Molecular Factors | References | Glossary
Related Pages: Placodes | Somite Development | Early Nervous | Early Heart | Abnormalities | Developmental Mechanisms | Epithelial Mesenchymal Transition | Molecular Development | References | Text only page | WWW Links |
Sweetman D, Wagstaff L, Cooper O, Weijer C, Munsterberg A. The migration of paraxial and lateral plate mesoderm cells emerging from the late primitive streak is controlled by different Wnt signals. BMC Dev Biol. 2008 Jun 9;8(1):63.
"...we use long-term video microscopy in chicken embryos to visualize the migration routes and movement behaviour of mesoderm progenitor cells as they emerge from the primitive streak (PS) between HH stages 7 and 10."
UNSW Embryology: Abnormalities | Sensory Placodes | Somite Development | Early Nervous System | 16 to 19Images - mesoderm and body cavities | 20 to 23Images - the yolk sac and gut formation | 24 to 25Images - mesoderm differentiation | 26 to 30Images - early placentation and fetal membranes | 31 to 34Images- placenta and cord
Embryo Images Online: Early Cell Populations and Establishment of Body Form | Neurulation (Week 4) | Form Changes, Gastrulation Continued and Neural Tube Closure (Weeks 4 and 5) | Neural Crest
Embryology Movies: Fate of the Somite (315Kb) | Vertebrae (366Kb) | Neurulation (468Kb) | Secondary Neurulation (561Kb)
Week 4 (Clinical Week 6)
Day |
Stage |
Event |
22 |
Stage 10 |
Neural Crest – differentiation at spinal cord level from day 22 until day 26 Neural – neural folds begin to fuse near the junction between brain and spinal cord, when Neural Crest cells are arising mainly from the neural ectoderm Neural Crest – trigeminal, facial, and postotic ganglia components visible (PMID: 17848161) Brain – rostral neural tube forms 3 primary brain vesicles (week 4) Respire – Week 4 - laryngotracheal groove forms on floor foregut. |
23 |
Heart – begins to beat in Humans by day 22-23, first functioning embryonic organ formed. Week 4 heart tube continues to elongate, curving to form S shape |
|
24 |
Stage 11 |
Thyroid – thyroid median endodermal thickening in the floor of pharynx Neural – rostral (or cephalic) neuropore closes within a few hours; closure is bidirectional, it takes place from the dorsal and terminal lips and may occur in two areas simultaneously. The two lips, however, behave differently. Optic ventricle appears |
25 |
Stage 12 |
Pituitary Week 4 – hypophysial pouch, Rathke’s pouch, diverticulum from roof GIT - Liver – septum transversum forming liver stroma and hepatic diverticulum forming hepatic trabeculae (PMID: 9407542) Neural – caudal neuropore takes a day to close (closure is approximately at future somitic pair 31/sacral vertebra 2) Neural – secondary neurulation begins Neural Crest – cardiac crest, neural crest from rhombomeres 6 and 7 that migrates to pharyngeal arch 3 and from there the truncus arteriosus (PMID: 17848161) |
26 |
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27 |
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28 |
Stage 13 |
Neural – the neural tube is normally completely closed, ventricular system now separated from amniotic fluid. Neural crest at spinal level is segregating, and spinal ganglia are in series with the somites. Spinal cord ventral roots beginning to develop. (PMID: 3354839) GIT - Liver – epithelial cord proliferation enmeshing stromal capillaries (PMID: 9407542) Sense - Smell – Crest comes from the nasal plates (PMID: 15604533) Skin – 4 weeks - simple ectoderm epithelium over mesenchyme Skin – 1-3 months ectoderm- germinative (basal) cell repeated division of generates stratified epithelium; mesoderm- differentiates into connective tissue and blood vessels |
Links: Textbooks | Online Textbooks | Search Textbooks | Reviews | Articles | Search PubMed | Glossary
Specific references can also be found on each notes page and Selected References 1999
Developmental Biology (6th ed) Gilbert, Scott F. Sunderland (MA): Sinauer Associates, Inc.; c2000. Figure 11.24. Implantation of the mammalian blastocyst into the uterus | Early Mammalian Development
Search NLM Online Textbooks "blastocyst implantation" : Developmental Biology | Endocrinology | Molecular Biology of the Cell | The Cell- A molecular Approach
Reviews
von Dassow M, Davidson LA. Variation and robustness of the mechanics of gastrulation: the role of tissue mechanical properties during morphogenesis. Birth Defects Res C Embryo Today. 2007 Dec;81(4):253-69.
Rohde LA, Heisenberg CP. Zebrafish gastrulation: cell movements, signals, and mechanisms. Int Rev Cytol. 2007;261:159-92.
Nance J, Lee JY, Goldstein B. Gastrulation in C. elegans. WormBook. 2005 Sep 26:1-13.
Kodaman PH, Taylor HS Hormonal regulation of implantation. Obstet Gynecol Clin North Am. 2004 Dec;31(4):745-66, ix.
Articles
Pézeron G, Mourrain P, Courty S, Ghislain J, Becker TS, Rosa FM, David NB. Live analysis of endodermal layer formation identifies random walk as a novel gastrulation movement. Curr Biol. 2008 Feb 26;18(4):276-81.
Nakaya Y, Sukowati EW, Wu Y, Sheng G. RhoA and microtubule dynamics control cell-basement membrane interaction in EMT during gastrulation. Nat Cell Biol. 2008 Jun 15.
Yao S, Xie L, Qian M, Yang H, Zhou L, Zhou Q, Yan F, Gou L, Wei Y, Zhao X, Mo X. Pnas4 is a novel regulator for convergence and extension during vertebrate gastrulation. FEBS Lett. 2008 Jun 25;582(15):2325-32.
Sweetman D, Wagstaff L, Cooper O, Weijer C, Munsterberg A. The migration of paraxial and lateral plate mesoderm cells emerging from the late primitive streak is controlled by different Wnt signals. BMC Dev Biol. 2008 Jun 9;8(1):63.
Güney M, Erdemoglu E, Oral B, Karahan N, Mungan T. Leukemia inhibitory factor (LIF) is immunohistochemically localized in tubal ectopic pregnancy. Acta Histochem. 2008 Feb 6;
Alisch RS, Jin P, Epstein M, Caspary T, Warren ST. Argonaute2 is essential for mammalian gastrulation and proper mesoderm formation. PLoS Genet. 2007 Dec 28;3(12):e227.
Hu W, Feng Z, Teresky AK, Levine AJ. p53 regulates maternal reproduction through LIF. Nature. 2007 Nov 29;450(7170):721-4.
Ralston A, Rossant J. How signaling promotes stem cell survival: trophoblast stem cells and shp2. Dev Cell. 2006 Mar;10(3):275-6.
Yang W, Klaman LD, Chen B, Araki T, Harada H, Thomas SM, George EL, Neel BG. An Shp2/SFK/Ras/Erk Signaling Pathway Controls Trophoblast Stem Cell Survival. Dev Cell. 2006 Mar;10(3):317-27.
Search Feb 2008 "blastocyst implantation" 7,336 reference articles of which 1,029 were reviews.
Search PubMed: term = week 3 embryonic human development | gastrulation | somite formation |
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