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rotates relative to the inner cytoplasm causes the *grey crescent* to form |
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MT driven gives rise to dorsal lip blastopore commit to invagination into the blastula initiates gastulation eventually forms the head endomesoderm and notochord can differentiate cells during gastulation |
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cell cycle goes from biphasic to quint phasic -blastopore ingresses into the embryo, causing a dorsal lip of the blastopore |
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acts a barrier to prevent vegatal cells from interacting with animal cells prematurely(induction) |
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found out that the vegatal cells release factors that tell the above cells (mesoderm) to become a specific thing |
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splitting or migrating of one sheet into two sheets |
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the expansion of one cell sheet over other cells Driven by Cell Division and Intercalation
Animal cap expand to cover the entire embryo • Mechanism includes: – Increase in cell division – Integration of cell deep layers into one |
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initiated on the future side of the dorsal side of the embryo-just below the grey crescent and across from the sperm entry |
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early cell movement during gastulation |
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cells ingress, then invaginate to form a slit, |
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cells changing their shape forming a bottleneck appearence
line the acheneron |
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•In salamanders--bottle cells appear to have an active role in the early movements of gastrulation •Dorsal marginal zone cells (which would normally give rise to the dorsal lip of the blastopore) were excised and placed on inner prospective endoderm tissue Bottle cells sank below the surface of the inner endoderm Bottle cells created a depression reminiscent of the early blastopore Bottle cells ability to invaginate into the deep endoderm is an innate property of the dorsal marginal zone cells |
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bottle cells bring the marginal cells in by involution -cells migrate along the basal surface of the bastocoel roof - |
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removal stops archenteron formation |
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– Involves filopodial extensions of involuted mesodermal cells that contact one another – Requires fibronectin |
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Differential Cell Cohesion |
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– Requires adhesion molecules that include paraxial protocadherin (somite-forming mesoderm) and axial protocadherin (notochord) • A dominant negative form of axial protocadherin (secreted and not membrane bound form) prevents convergent extension |
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– Increase observed cell/tissues undergoing convergent extension • Regulate contraction of actin filaments |
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oocyte has animal vegetal polarity
A-V provides information about which cells will form which germ layer |
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vegetal cells(2 functions) |
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differentiate into endoderm induce cells above them to become mesoderm |
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dorsal is opposite sperm entry |
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movement on invoulting tissue(head mesoderm)during gastulation |
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process forming dorsal-ventral and anterior-posterior |
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dorsal blastopore lip -primary embryonic induction -Induce ventral tissues to change fate to form neural tube and dorsal mesoderm (somites) -Organize through initial inductive events the embryos anterior-posterior and dorsa-ventral axes |
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ligation experiment
conclusion from experiment: early amphibian muclei were indentical each cell was capable of developing a complete organism |
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fate depends on their location in the embryo(cells are uncommitted) |
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their fate was already determined(epidermal and neural) |
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primary embryonic induction |
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Numerous inductions during embryonic development— key induction dorsal lip cells induce dorsal axis and neural tube |
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induction forms the organizer |
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recieves signals from the endoderm(the organizer does) -dorsal most vegetal cells can induce the organizer |
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dorsal most vegetal cells induce animal cells to be come dorsal mesoderm |
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Transplantation and Recombination Experiments (1986)—Demonstrated the existence of the Nieuwkoop Center |
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Β-Catenin is localized on Dorsal side •Duel Function: Transcription factor and transmembrane protein •Location:Prior to fertilization-- Β-Catenin located throughout oocyte Shortly after fertilization--Β-Catenin accumulates dorsal region during early cleavage |
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Nieuwkoop Center secretes TGF- Β Family paracrine factors of which induces expression of Smad2/4 – Vg1 belongs to TGF-Β family – Other TGF-Β Family include activin, and Derriere |
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Activation of Organizer by binding of transcription factors Smad2/4 and Siamois |
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a DNA binding protein -activated by Nieuwkoop center activates the migration properties (involution and convergent extension) of the dorsal blastopore lip cells – autonomously determines the dorsal mesodermal fates of those cells expressing it – enables the goosecoid-expressing cells to recruit neighboring cells into the dorsal axis. |
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Β-catenin acts synergistically with VegT and Vg1 to activate transcription of Xnr (Nodal family) Vg1 and Xnr = TGF-B Family |
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To initiate the movements of gastrulation • To become dorsal mesoderm • To dorsalize the surrounding mesoderm into lateral mesoderm (when it would otherwise form ventral mesoderm) • To dorsalize the ectoderm into neural ectoderm • To cause the neural plate (the induced neural ectoderm) to become the neural tube
Inhibits Epidermal Inducer, Thus Neural Tissue is Formed |
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Induction of Neural Ectoderm and Dorsal Mesoderm • Ectoderm is induced to become epidermal tissue by binding Bone Morphogeneic Proteins (BMPs; ex BMP4) – BMPs are powerful ventralizing factors • Form dorsalateral muscle, ventral mesenchyme (blood), ventral lateral plate (kidney), and epidermal ectoderm (skin) • Three major BMP inhibitors include Noggin, Chordin, Follistatin – Promotes Dorsal Endoderm and Neural Ectoderm to form |
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Organizer and Anterior-Posterior Axis and strucure |
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In Xenopus (and other vertebrates), the formation of the anteriorposterior axis follows the formation of the dorsal-ventral axis. • Once the dorsal portion of the embryo is established, the movement of the involuting mesoderm establishes the anterior-posterior axis • The mesoderm that migrates first through the dorsal blastopore lip gives rise to the anterior structures take place at the dorsal blastopore lip |
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Regional specific induction |
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•Cells of the organizer to enter the embryo first induce the formation of brain and head •Cells entering from the dorsal lip of later-stage embryos induce the cells to become spinal cord and tail |
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Organizer Function and A-P Axis Specification in Xenopus |
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A. Head and Brain Formation: – Wnt and BMP inhibitors (1 and 2) in the anterior of the organizer (pharyngeal mesendoderm) allow the induction of head structures by blocking BMP 4 and XWnt8 • B. Neural Ectoderm and Trunk Inducer: (1) – BMP inhibitors from organizer tissue block the formation of epidermis whereas Wnt induces neural tissue to form • C. Caudal Inducer: (3) – Gradient of caudalizing factors (eFGF, retinoic acid, and/or Wnt3a) specify regions of neural tube through expression of Hox genes |
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Left-Right Axis Formation |
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Two separate molecular mechanisms used to form L-R axis – Regulated by two factors: paracrine Nodal (Xnr1) and transcription factor Pitx2 • Nodal gene (Xnr1) only expressed on the lateral plate mesoderm on left side of embryo – Cilia may also play a role in establishing L-R axis • Cells in organizer have cilia that beat clockwise flow of fluid—possible sweeping Xnr1 predominately to the left and thus activating Pitx2 – Pitx2—controls position of heart and gut |
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