GASTRULATION IN AMPHIBIANS

DEFINITION:
Gastrulation is the process during embryonic development that changes the embryo
from a blastula with a single layer of cells to a gastrula containing multiple layers
of cells.
DESCRIPTION:
The layers created by gastrulation become germ layers, or special tissues that give
rise to specific parts of the organism. These germ layers always give rise to the
same types of tissues, even in very different animals.
 The endoderm will give rise to the gut and associated organs.
 The ectoderm is the outermost layer, and will create the skin and the nervous
system.
 Between them lies the mesoderm, which will created the connective tissues
and musculature in most organisms.

AMPHIBIAN GASTRULATION

Amphibian Gastrulation:
Quick Overview
1.Gastrulation begins at the future dorsal side, below the equator (marginal zone)
when prospective endodermal cells (bottle cells) invaginate and form a slit
(blastopore). The bottle cells line the early archenteron.

2. The marginal zone cells undergo involution at the dorsal lip of the blastopore.

3. After the bottle cells, the next cells that enter the embryo form the prechordal
plate (head mesoderm), and the mesoderm of the notochord.

4. Simultaneously, the animal cells undergo epiboly, completely surrounding the

surface of the embryo until a ventral lip of the blastopore forms.

5. The last endodermal cells on the surface appear as the yolk plug.

6. The yolk plug is eventually covered by ectoderm

AMPHIBIAN GASTRULATION:

The study of amphibian gastrulation is both one of the oldest and one of the newest
areas of experimental embryology. The study of amphibian gastrulation has been
complicated by the fact that there is no single way amphibians gastrulate. Different
species employ different means toward the same goal). In recent years, the most
intensive investigations have focused on the frog Xenopus laevis, so we will
concentrate on its mode of gastrulation.

Fig: frog gastrula

 THE FATE MAP OF XENOPUS

Amphibian blastulae are faced with the same tasks as the invertebrate blastulae, 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. The movements whereby
this is accomplished can be visualized by the technique of vital dye staining.
Fate mapping by Løvtrup (1975; Landstrom and Løvtrup 1979) and by Keller
(1975,1976) has shown that cells of the Xenopus blastula have different fates
depending on whether they are located in the deep or the superficial layers of the
embryo .In Xenopus, the mesodermal precursors exist mostly in the deep layer of
cells, while the ectoderm and endoderm arise from the superficial layer on the
surface of the embryo. Most of the precursors for the notochord and other
mesodermal tissues are located beneath the surface in the equatorial (marginal)
region of the embryo. In urodeles (salamanders such as Triturus and Ambystoma)
and in some frogs other than Xenopus, many more of the notochord and mesoderm
precursors are among the surface .

Fig : Fate maps of the blastula of the frog Xenopus laevis: (A) exterior; (B) interior.
As we have seen, the unfertilized egg has a polarity along the animal-vegetal axis.
the germ layers can be mapped onto the oocyte even before fertilization.
 The surface of the animal hemisphere will become the cells of the ectoderm
(skin and nerves).
  the vegetal hemisphere surface will form the cells of the gut and associated
organs (endoderm), and the mesodermal cells will form from the internal
cytoplasm around the equator.
FACTORS CONTROLLING GASTRULATION IN AMPHIBIANS
This general fate map is thought to be imposed upon the egg by the transcription
factor VegT and the paracrine factor Vg1. The mRNAs for these proteins are
located in the cortex of the vegetal hemisphere of Xenopus oocytes, and they are
apportioned to the vegetal cells during cleavage By using antisense oligonucleotides,
Zhang and colleagues (1998) were able to deplete maternal VegT protein in early
embryos. The resulting embryos lacked the normal fate map. The animal third of the
embryo produced only ventral epidermis, while the marginal cells (which normally
produced mesoderm) generated epidermal and neural tissue. The vegetal third
(which usually produces endoderm) produced a mixture of ectoderm and mesoderm
demonstrated that embryos that lacked functional Vg1 lacked endoderm and dorsal
mesoderm.

Fig : The fates of the three regions of the Xenopus blastula are altered by the
depletion of VegT

CELL MOVEMENTS DURING AMPHIBIAN GASTRULATION

Gastrulation in frog embryos is initiated on the future dorsal side of the embryo, just
below the equator in the region of the gray crescent .Here, the cells invaginate to
form a slitlike blastopore. These cells change their shape dramatically. The main
body of each cell is displaced toward the inside of the embryo while the cell
maintains contact with the outside surface by way of a slender neck. These bottle
cells line the archenteron as it forms. Thus, as in the gastrulating sea urchin, an
invagination of cells initiates archenteron formation. However, unlike gastrulation
in sea urchins, gastrulation in the frog begins not at the most vegetal region, but in
the marginal zone: the zone surrounding the equator of the blastula, where the
animal and vegetal hemispheres meet. Here the endodermal cells are not as large or
as yolky as the most vegetal blastomeres.

Fig: Cell movements during frog gastrulation

The next phase of gastrulation involves the involution of the marginal zone cells
while the animal cells undergo epiboly and converge at the blastopore .
When the migrating marginal cells reach the dorsal lip of the blastopore, they turn
inward and travel along the inner surface of the outer animal hemisphere cells. Thus,
the cells constituting the lip of the blastopore are constantly changing. The first cells
to compose the dorsal blastopore lip are the bottle cells that invaginated to form the
leading edge of the archenteron. These cells later become the pharyngeal cells of the
foregut.

Fig : Epiboly of the ectoderm

THE MIDBLASTULA TRANSITION: PREPARING FOR

GASTRULATION

Now that we have seen an overview of amphibian gastrulation, we can look more
deeply into its mechanisms. The first precondition for gastrulation is the activation
of the genome. In Xenopus, the nuclear genes are not transcribed until late in the
twelfth cell cycle .At that time, different genes begin to be transcribed in different
cells, and the blastomeres acquire the capacity to become motile. This dramatic
change is called the midblastula transition.

POSITIONING THE BLASTOPORE

The vegetal cells are critical in determining the location of the blastopore, as is the
point of sperm entry. The microtubules of the sperm direct cytoplasmic movements
that empower the vegetal cells opposite the point of sperm entry to induce the
blastopore in the mesoderm above them. This region of cells opposite the point of
sperm entry will form the blastopore and become the dorsal portion of the body.
Figure 10.11: Transplantation experiments on 64-cell amphibian embryos
demonstrating that the vegetal cells underlying the prospective dorsal blastopore
lip region are responsible for causing the initiation of gastrulation. (A) Rescue of
irradiated embryos by transplanting the dorsal vegetal blastomeres of a normal
embryo into a cavity made by the removal of a similar number of vegetal cells. An
irradiated zygote without this transplant fails to undergo normal gastrulation. (B)
Formation of a new gastrulation site and body axis by the transplantation of the
most dorsal vegetal cells of one embryo into the ventralmost vegetal region of
another embryo.

INVAGINATION AND INVOLUTION

Amphibian gastrulation is first visible when a group of marginal endoderm cells on

the dorsal surface of the blastula sinks into the embryo. The outer (apical) surfaces
of these cells contract dramatically, while their inner (basal) ends expand. The
apical-basal length of these cells greatly increases to yield the characteristic “bottle”
shape. In salamanders, these bottle cells appear to have an active role in the early
movements of gastrulation. Johannes Holtfreter (1943,1944) found that bottle cells
from early salamander gastrulae could attach to glass coverslips and lead the
movement of those cells attached to them. Even more convincing were Holtfreter's
recombination experiments. When 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, they formed bottle cells and sank below the
surface of the inner endoderm (Figure 10.12). Moreover, as they sank, they created
a depression reminiscent of the early blastopore. Thus, Holtfreter claimed that the
ability to invaginate into the deep endoderm is an innate property of the dorsal
marginal zone cells.

Figure : A graft of cells from the dorsal marginal zone of a salamander embryo
sinks into a layer of endodermal cells and forms a blastopore-like groove.
CONCLUSION:
The layers created by gastrulation become germ layers, or special tissues that give
rise to specific parts of the organism. These germ layers always give rise to the
same types of tissues, even in very different animals. The endoderm will give rise
to the gut and associated organs. The ectoderm is the outermost layer, and will
create the skin and the nervous system. Between them lies the mesoderm, which
will created the connective tissues and musculature in most organisms.
REFERENCES:
 https://www.ncbi.nlm.nih.gov/books/NBK10113/figure/A2284/?report=obje
ctonly
 http://www.yourarticlelibrary.com/notes/useful-notes-on-gastrulation-in-
frog-biology/5151
 https://biologydictionary.net/gastrulation/
 https://www.biology-pages.info/F/FrogEmbryology.html
 https://www.researchgate.net/figure/Diagrams-of-hypothetical-
amphibian-gastrulation-movements-A-B-show-that-
involution_fig9_8570029
 CONTENTS

 Gastrulation in amphibians

 Quick overview

 Fatemap of Xenopus

 Factors controlling gastrulation in amphibians

 Cell movements during amphibian gastrulation

 Midblastula transitions

 Positioning the blastopore

 Invaginations and involution

TOPIC: GASTRULATION IN AMPHIBIANS

SUBMITTED BY : SAMREEN

ROLL NO :

SUBMITTED TO : Dr. SAIMA JABEEN

SEMESTER : VI

COURSE TITLE : DEVELOPMENTAL BIOLOGY

COURSE CODE : Z00-

DATE OF SUBMISSION: 21ST FEB, 2020

DEPARTMENT OF ZOOLOGY
UNIVERSITY OF GUJRAT