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  • Lecture 16 Fate Mapping and Gastrulation in Sea Urchin

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    IQra Kanwal
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    1. During sea urchin development, there is a cascade of interactions between blastomeres where micromeres induce veg2 cells and veg2 cells induce veg1 cells. 2. β-catenin, activated by the Wnt pathway, is responsible for specifying the micromeres and their inductive ability. It accumulates in nuclei of cells fated to become endoderm and mesoderm. 3. During gastrulation, micromeres ingress into the blastocoel by losing affinity for the hyaline layer and their neighbors and gaining affinity for proteins in the blastocoel lining. They migrate and arrange along the animal-vegetal axis, forming clusters that will become

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    1. During sea urchin development, there is a cascade of interactions between blastomeres where micromeres induce veg2 cells and veg2 cells induce veg1 cells. 2. β-catenin, activated by the Wnt pathway, is responsible for specifying the micromeres and their inductive ability. It accumulates in nuclei of cells fated to become endoderm and mesoderm. 3. During gastrulation, micromeres ingress into the blastocoel by losing affinity for the hyaline layer and their neighbors and gaining affinity for proteins in the blastocoel lining. They migrate and arrange along the animal-vegetal axis, forming clusters that will become

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    436 views23 pages

    Lecture 16 Fate Mapping and Gastrulation in Sea Urchin

    Uploaded by

    IQra Kanwal
    1. During sea urchin development, there is a cascade of interactions between blastomeres where micromeres induce veg2 cells and veg2 cells induce veg1 cells. 2. β-catenin, activated by the Wnt pathway, is responsible for specifying the micromeres and their inductive ability. It accumulates in nuclei of cells fated to become endoderm and mesoderm. 3. During gastrulation, micromeres ingress into the blastocoel by losing affinity for the hyaline layer and their neighbors and gaining affinity for proteins in the blastocoel lining. They migrate and arrange along the animal-vegetal axis, forming clusters that will become

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    of 23

    Lecture 16

    Gastrulation in Sea Urchin


    Interaction of blastomeres
    • In a normal embryo
    • Micromeres regulate specification of
    veg2 cells
    • Veg 2 cells specify veg1 layer.

    • In the absence of veg2 layer


    • veg1 cells produce endoderm
    • But the endoderm does not specify
    into foregut, midgut, or hindgut.
    Interaction of blastomeres
    • So, there is a cascade of interaction in which

    • The vegetal pole micromeres induce the cells above them to


    become the veg2 cells

    • The veg2 cells induce the cells above them to assume the veg1
    fates
    Interaction of blastomeres
    • It is hypothesized that

    • Signals from the micromeres induce a post-translational


    modification in some factor in the veg2 layer.

    • Signals from the veg2 layer probably modify some protein


    (transcription factor) in the veg1 layer.
    Interaction of blastomeres
    • The molecule responsible for specifying the micromeres appears to
    be β-catenin

    • β-catenin is a transcription factor


    • Activated by the Wnt pathway
    Interaction of blastomeres

    • Several pieces of evidence suggest this role of β-catenin .

    1. During normal sea urchin development


    • β-catenin accumulates in the nuclei of cells fated to
    become endoderm and mesoderm
    Interaction of blastomeres
    2. This accumulation is responsible for specifying the
    vegetal half of the embryo.

    • Treatment of sea urchin embryos with lithium chloride

    • Causes the accumulation of β-catenin in all


    embryonic cells
    • Transforms the presumptive ectoderm into
    endoderm
    Interaction of
    blastomeres
    2. This accumulation is responsible for specifying
    the vegetal half of the embryo.

    • Inhibition of β-catenin accumulation in the


    vegetal cell nuclei
    • Prevent the formation of endoderm and
    mesoderm
    Interaction of blastomeres

    3. β-catenin is essential for giving the


    micromeres their inductive ability

    • Micromeres from embryos without β-catenin


    in their nucleus

    • Do not induce the animal hemisphere cells to


    form endoderm
    Axis specification in sea urchin
    • In the sea urchin

    • Anterior-posterior axis is specified before fertilization


    • The animal-vegetal axis gives rise to anterior-posterior
    axis

    • Dorsal-ventral axis is specified after the 8-cell stage


    (3divisions)

    • The left-right axes are specified after fertilization.


    Sea Urchin Gastrulation
    • The late sea urchin blastula consists of
    • single layer of about 1000 cells
    • a hollow ball
    • somewhat flattened at the vegetal end

    • The blastomeres, derived from different regions of the


    zygote have
    • different sizes
    • different properties

    • Blastula develops into pluteus larva through gastrulation


    Ingression of primary mesenchyme
    • Shortly after the blastula hatches from its fertilization
    envelope

    • the vegetal side of the spherical blastula begins to thicken


    and flatten
    Filopodia formation
    • At the center of this flat vegetal plate,
    • a cluster of small cells form contractile long, thin
    processes----- filopodia from their inner surfaces.

    • The cells then dissociate from the epithelial monolayer


    and ingress into the blastocoel
    Ingression of primary mesenchyme

    • These cells, derived from the micromeres ----


    ---- primary mesenchyme

    • They will form the larval skeleton -------


    skeletogenic mesenchyme
    Spicule formation
    • At first the cells appear to move randomly along the
    inner blastocoel surface
    • Actively make and break filopodial connections to the
    wall of the blastocoel

    • Eventually, they become localized within the future


    ventrolateral region of the blastocoel.

    • Here they fuse into syncytial cables, which will form


    the axis of the calcium carbonate spicules of the larval
    skeleton.
    Mechanism of ingression of primary mesenchymes
    • The ingression of the micromere descendants into the blastocoel is
    caused by
    • loss of affinity to
    • their neighbors
    • hyaline membrane
    • acquiring a strong affinity for
    • a group of proteins present in the lining of the blastocoel

    • Originally, all the cells of the blastula are connected on their outer
    surface to the hyaline layer and on their inner surface to a basal
    lamina secreted by the cells.
    Mechanism of ingression of primary mesenchymes
    • The primary mesenchyme precursors
    • Lose their affinity for
    • the hyaline layer
    • their neighbors
    • Increase their affinity hundredfold for
    • basal lamina
    • extracellular matrix (such as fibronectin)

    • There is a changes in cell surface molecules during this time.


    Interaction of remaining mesomeres and macromeres

    • Ectoderm (descendants of the mesomeres) and endoderm cells


    (descendants of macromeres) bind tightly to
    • one another
    • to the hyaline layer

    • Adhere only loosely


    • to the basal lamina
    Micromere Migration
    • However, the micromere pattern changes during gastrulation
    • This change in affinity causes the micromeres
    • to loss their connections with
    • external hyaline layer
    • their neighboring cells

    • They are then attracted by the basal lamina


    • Migrate up into the blastocoel
    Micromere Migration
    • Inside the blastocoel, primary mesenchyme cells
    • migrate along the extracellular matrix of the blastocoel wall
    • extend their filopodia in front of them

    • Several proteins (fibronectin and a particular sulfated


    glycoprotein) are necessary to initiate and maintain this
    migration
    Micromere arrangement
    • The primary mesenchyme cells arrange
    themselves in a ring at a specific position
    along the animal-vegetal axis.

    • At two sites near the future ventral side of the


    larva
    • many primary mesenchyme cells form
    cluster
    • fuse with each other
    • Initiate spicule formation
    Cues for Micromere Migration
    • This positional information is provided by
    • future ectodermal cells
    • basal laminae

    • Skeleton-forming mesenchyme cells contain


    • extremely fine (0.3-μm diameter) filopodia
    • To explore and sense dorsal-ventral and animal-vegetal patterning
    cues from the ectoderm

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