Gastrulation: Highly integrated cell and tissue
movements whereby blastula cells are dramatically
rearranged
-3 primary germ layers are established
-basic body plan is established (rudimentary primary and
body axes)
-cells brought to new positions, for inductive interactions
(neurulation and organogenesis)
*Fate has been specified but potency not fully
determined
*Polarity of egg (Vegetal-Animal pole) when established
as the zygote is formed determine anterior and posterior
axes
*Cells that use to be together at the end of the cleavage
will be rearranged
Inductive interactions: cells have autonomous (cell will
always give rise to its fate whether where it is located)
and conditional (dependent to the signal of surrounding
cells) development
Organs do not form on their own, they need signals
>Ectoderm: Epidermis, brain, NS
>Endoderm: Embryonic gut
>Mesoderm: Notocord
Embryonic Cell Movement: heart of gastrulation
-goal is to form the three germ layers
Embryonic Cellular Movements:
>Invagination- movement of a group of cells (epithelial)
inward: inpocketing [opposite: evagination- lumen
formed by basal surface]
>Involution- rolling movement, free to move further up
underneath the exterior tissue (by bulk)
>Ingression- individual cells move away from epithelial
sheets migrating inwards, usually mesenchymal cells
(with cytoplasmic extension) ex. neural crest cell HOW:
-alter cell architecture
-alter program of motility
-alter adhesiveness (lose it, change in micro-
tubule alignment)
*SPREADING OF GROUP OF CELLS
>Epiboly- increase in cell surface area, change in cell
shape; resulting to thinning of cells so that it can cover
another area (monolayer)
>Intercalation- 2 or more rows of cells, moving to one
another, insertion of cells to junction of other layer,
increasing surface area of tissue, cell rearrangement
>Convergent extension- specialized intercalation,
however it has a specific direction to follow. Increase in
surface area of tissue. Intercalating perpendicular to the
axis of extension
>Delamination- separation of two or more layers.
Common in hypoblast, epiblast layers
Consider:
-Unit of migration (individual or sheet?)
-Intrinsic/Extrinsic factor in movement
-Spreading: movement of tissue
-Change in shape and motility (columnar, bottle,
mesenchymal)
MODEL: Sea Urchin Gastrulation
(colors: fate map)
Mesomeres- animal pole: conditional
Macromeres- vegetal cells: conditional
Micromeres: autonomous specification (skelotegenic
cells) produce skeletal spicules
Specifies veg2 cells that specify veg1 layer-
produce endoderm
Proteins (in cell specification): Beta catenin
(transcription factor)- operates by Wnt pathway,
molecule responsible for specifying micromeres,
inducing other areas to form structures
>no involution only invagination (presumptive
endoderm: vegetal region/plate) due to changes in cell
shape to form archenteron for elongation
>Primary Mesenchymal Cells: ingression
>Vegetal plate: primary invagination (result from
changes in shape of cells in vplate) to produce
archenteron (primitive gut)
>Secondary invagination- elongation of archenteron
across blastocoel: attches near animal pole of embryo
(filopodia extended by secondary mesenchyme cells
locates at the tip of archenteron involved)
>Secondary Mesenchymal Cells: from tip of invaginating
archenteron- changes in cells in the tip performing
ingression: drags elongating archenteron to contact with
ectodermal cells
>Apical constriction and invagination- vegetal plate
endoderm cells change their shape from a cuboidal to
columnar to flask-shape (intercalates and extends
elongating the archenteron tube upward towards roof of
blastocoel cavity, bend to left, fuse and eventually forms
future oral opening)
Amphibian: Tasks of gastrulation
• to bring inside the embryo those areas destined
to form the endodermal organs,
• to surround the embryo with cells capable of
forming the ectoderm, and
• to place the mesodermal cells in the proper
positions between them
Fate map of Xenopus gastrula
*Imposed upon the egg by transcription factor VegT
(autonomic: endoderm) and paracrine factor Vg1
1.) different gastrula areas show distinct cellular
behaviors
• Animal cap
– Engage in epiboly
• Non-involuting marginal zone
– Undergoes epiboly
• Involuting marginal zone
– Forms upper margin of blastopore
– Superficial IMZ
• Forms the archenteron roof
– Deep IMZ
• Gives rise to much of the mesoderm
• Deep Zone
– Ring of cells between DIMZ & large vegetal cells
– Migrate up wall & roof of blastocoel; Leads IMZ
• Bottle cells
– Vegetal border of IMZ
• Vegetal base
– Large yolky cells extending from bottom of
blastocoel to outside of vegetal hemisphere
– Displaced ventrally during gastrulation
– Forms archenteron floor
2. Bottle cells generate initial depression of blastopore
Frog Gastrulation begins in marginal zone ( surround
equator of blastula, animal and vegetal hemispheres
meet. Blastopore lip formed by bottle cells, changing cell
shape: apical constriction creates local invagination
*Jeff Hardin and Ray Keller: culture explants in surgically
altered embryos :isolated bottle cells constrict at apical
surface but assume characteristic bottle shape only with
other cells surrounding
*Xenopus (RE 1981) bottle cells are not necessary for
continued gastrulation; role is limited to generating
initial depression of the blastopore
–Removal of bottle cells: embryo healed w/in
30min, gastrulated and develop normally except that the
archenteron is truncated at cephalic end
3. Involution of marginal zone cells: next phase of
gastrulation; occurs while animal cells undergo epiboly
and converge at blastopore
(RE Keller 1981) Replace involuting marginal zone layers
with other gastrula tissues (both superficial layer and
deep layer w/ animal cap: grafted animal cap patches
balked at blastopore lip and did not move in) (superficial
layer with animal cap: involuted and became part of
archenteron roof)
Bottle cells to pharyngeal cells of foregut, DB become
prechordal plat, next cells involuting into the embryo
through DBL- chordamesoderm cells: form notochord)
4. Convergent Extension is especially strong in Dorsal
Marginal Zone- participated by NIMZ and IMZ
*Keller et al., 1985: Radical intercalation- perpendicular
to embryo surface, Mediolateral intercalation: parallel to
embryo surface
5. The Animal Cap and the non-ivoluting marginal zone
undergo epiboly- thinning, cell division, rearrangement:
decrease cell layers in animal cap from 2 to 1: intercalate
-Events involve during epiboly: radical intercalation in
deep layer
 Mechanisms on Xenopus: 2 signaling pathways-
Planar (major role in embryos) sent by
blastopore lip through ectoderm plane (express region-
specific marker genes) and Vertical signal sent to
overlying ectoderm: the prospective neural plate
Neuralizing mesoderm signals
3. Neural induction is a multistep process (associated
with region specific expression of genes)
*Ectoderm devt depends on proximity of inducing tissue
and preparedness of responding tissue
-Dorsal ectoderm: Neural plate
-Ventral ectoderm: Epidermis; expresses Epi1 at
midneurula stage

Neural Induction
-formation of neural tube in association w/ an underlying
notochord, laterally adjacent somites, and other
mesodermal structures EMBRYONIC AXIS: stereotypical
arrangement; coordinated dev’t by inductive
interactions (blastopore lip)
1. Dorsal blastopore lip organizes formation of entire
embryo (Spemann experiments)
*graft of dbp developed accdg to fate: notochord
*graft dorsalized the host’s mesoderm: kidneys
*graft acted as neural inducer; host ectoderm formed as
neural plate
2. Neural induction shows regional specificity:
*Experiment by Spemann Blastopore Graft (early
gastrula: complete head no tail): Early-involuting
mesoderm tended to induce anterior axial structures;
Later-involuting mesoderm induce trunk and tail
*Otto Mangold Experiment: Neural plate form early
neurula exposing chordamesoderm (notochord and
adjacent somites) Chordamesoderm was isolated and
divided into 4 Differentiation of Neural Tube (give rise to glial cells and
1st quarter: induced parts that lie in front of brain neurons)
2nd quarter: head+brain+eyes+ear vesicles -consist of neuroepithelial cells
3rd quarter: hindbrain+spinal cord+ musculature -lies on a basement membrane, single layer
4th quarter: spinal cord+ somites + kidney + tail -neural epithelial cells: germinal epithelium
-blastopore lip at diff stages and corresponding areas of -neural stem cells: divide assymetrically
chordamesoderm from late gastrula induce diff sets of Stem cells
dorsal structures. Committed progenitor cell (PMC ingression)
Spinal cord: distinction between 3 layers (meninges,
white and gray matter) differ in bulge formation from
brain
*1st stage: cells break away from lumen, migrate towards
external limiting membrane: outside (these cells form
mantle that eventually becomes gray matter) maturation
are noted by axons and dendrites (further maturation
sheathing of schwann cells: formation of marginal layer:
white matter)
-white matter>grey matter>Ependymal cells (csf
formation)
-when differentiating nucleus go to basal region for
development

*Formation of dorsal and ventral ridges

Alar plate: Dorsal- roof plate
Basal plate: Ventral- floor plate
*Notochord for dorso-ventral patterning
:Add notochord= floor plate- devt dorsal neurons

Meten cerebrum
Mylen medulla
Origin of PNS Cells
Neural Crest Cells: varied fates
-determined by neural plate/ectoderm and epidermal
ectoderm are adjacent to each other
-boundary of neural plate and epidermis
-arise from inductive interaction of epidermis and neural
epidermis
-arise from neural plate cells and epidermal cells
-acquire properties that can be distinguished from
neighboring cells: inductive influence of
chordamesoderm (intermediate signal) or juxtaposition
hypothesis c