Frog Embryology

The Egg
The frog egg is a huge cell; its volume is over 1.6 million times larger than a normal
frog cell. During embryonic development, the egg will be converted into a tadpole
containing millions of cells but containing the same amount of organic matter.
The upper hemisphere of the egg the animal pole is dark.
The lower hemisphere the vegetal pole is light.

When deposited in the water and ready for fertilization, the haploid egg is at
metaphase of meiosis II

Fertilization
Entrance of the sperm initiates a sequence of events:
Meiosis II is completed.
The cytoplasm of the egg rotates about 30
degrees relative to the poles.

In some amphibians (including Xenopus), this

is revealed by the appearance of a light-colored
band, the gray crescent.

The gray crescent forms opposite the point

where the sperm entered.

It foretells the future pattern of the animal: its dorsal (D) and ventral (V)
surfaces; its anterior (A) and posterior (P); its left and right sides.

The haploid sperm and egg nuclei fuse to form the diploid zygote nucleus.

Cleavage
The zygote nucleus undergoes a series of mitoses, with the resulting daughter nuclei
becoming partitioned off, by cytokinesis, in
separate, and ever-smaller, cells. The first
cleavage occurs shortly after the zygote
nucleus forms.
A furrow appears that runs
longitudinally through the poles of
the egg, passing through the point at
which the sperm entered and
bisecting the gray crescent.

This divides the egg into two halves forming the 2-cell stage

The second cleavage forms the 4-cell stage. The cleavage furrow again runs through
the poles but at right angles to the first furrow.
The furrow in the third cleavage runs horizontally but in a plane
closer to the animal than to the vegetal pole. It produces the 8-cell
stage.
The next few cleavages also proceed in synchrony, producing a 16cell and then a 32-cell embryo.
However, as cleavage continues, the cells in the animal pole begin dividing more
rapidly than those in the vegetal pole and thus become smaller and more numerous.
By the next day, continued cleavage has produced a hollow ball of thousands of cells
called the blastula. A fluid-filled cavity, the blastocoel, forms within it.
During this entire process
there has been no growth of the
embryo. In fact, because the cells
of the blastula are so small, the
blastula looks just like the original
egg to the unaided eye.
Not until the blastula contains
some 4,000 cells is there
any transcription of zygote genes. All of the activities up to now have been run
by gene products (mRNA and proteins) deposited by the mother when she
formed the egg.

Gastrulation
The start of gastrulation is marked by the pushing inward ("invagination") of cells in
the region of the embryo once occupied by the middle of the gray crescent. This
produces:
an opening (the blastopore) that will be the future
anus
a cluster of cells that develops into the Spemann
organizer (named after one of the German
embryologists who discovered its remarkable
inductive properties).
As gastrulation continues, three distinct "germ layers" are
formed:
ectoderm
mesoderm

endoderm

Each of these will have special roles to play in building the complete animal. Some
are listed in the table.
Germ-layer origin of various body tissues
Ectoderm

Mesoderm

Endoderm

skin

notochord inner lining of gut, liver, pancreas

brain

muscles

inner lining of lungs

spinal cord

blood

all other neurons bone

sense receptors

thyroid and parathyroid glands

sex organs thymus

The Spemann

inner lining of bladder

organizer (mostly mesoderm) will:

develop
into the notochord, which is the precursor of the backbone;
induce the ectoderm lying above it to begin to form neural tissue instead of
skin.
o

This ectoderm grows up into two longitudinal folds, forming the neural
folds stage.

In time the lips of the folds fuse to form the neural tube.

The neural tube eventually develops into the brain and spinal cord.

Differentiation
Although the various layers of cells in the frog gastrula have definite and different
fates in store for them, these are not readily apparent in their structure. Only by
probing for different patterns of gene expression (e.g., looking for tissue-specific
proteins) can their differences be detected.
In due course, however, the cells of the embryo take on the specialized structures and
functions that they have in the tadpole, forming neurons, blood cells, muscle
cells, epithelial cells, etc., etc.

Growth
At the time the tadpole hatches, it is a fully-formed organism. However, it has no
more organic matter in it than the original frog egg had. Once able to feed, however,
the tadpole can grow. It gains additional molecules with which it can increase the
number of cells that make up its various tissues.