Early Development & Formation
Patterns of cleavage
● The cell, manifestly, entertains a very different opinion.”
● Indeed, different organisms undergo cleavage in distinctly different ways, and the mechanisms
for these differences remain at the frontier of cell and developmental biology.
● Cells in the cleavage-stage cells are called blastomeres.
● In most species mammals being the chief exception, both the initial rate of cell division and
the placement of the blastomeres with respect to one another are under the control of
proteins and mRNAs stored in the oocyte.
● Only later do the rates of cell division and the placement of cells come under the control of
the newly formed organism’s own genome.
● During the initial phase of development, when cleavage rhythms are controlled by maternal
factors, cytoplasmic volume does not increase.
● Rather, the zygote cytoplasm is divided into increasingly smaller cells—first in half, then
quarters, then eighths, and so forth.
● Cleavage occurs very rapidly in most invertebrates, probably as an adaptation to generate a
large number of cells quickly and to restore the somatic ratio of nuclear volume to cytoplasmic
volume.
● The embryo often accomplishes this by abolishing the gap periods of the cell cycle the G1 and
G2 phases, when growth can occur.
● A frog egg, for example, can divide into 37,000 cells in just 43 hours.
● The pattern of embryonic cleavage peculiar to a species is determined by two major
parameters:
a. The amount and distribution of yolk protein within the cytoplasm, which determines
where cleavage can occur and the relative sizes of the blastomeres; and
b. Factors in the egg cytoplasm that influence the angle of the mitotic spindle and the
timing of its formation.
● In general, yolk inhibits cleavage.
● When one pole of the egg is relatively yolk-free, cellular divisions occur there at a faster rate
than at the opposite pole.
● The yolk-rich pole is referred to as the vegetal pole; the yolk concentration in the animal pole
is relatively low.
● The zygote nucleus is frequently displaced toward the animal pole.
● At one extreme are the eggs of sea urchins, mammals, and snails.
● These eggs have sparse, equally distributed yolk and are thus isolecithal Greek, “equal yolk”.
In these species, cleavage is holoblastic Greek holos, “complete”, meaning that the cleavage
furrow extends through the entire egg.
● With little yolk, these embryos must have some other way of obtaining food.
● Most will generate a voracious larval form, while mammals will obtain their nutrition from the
maternal placenta.
● At the other extreme are the eggs of insects, fish, reptiles, and birds.
● Most of their cell volumes are made up of yolk. The yolk must be sufficient to nourish these
animals throughout embryonic development.
● Zygotes containing large accumulations of yolk undergo meroblastic cleavage Greek meros,
“part”, wherein only a portion of the cytoplasm is cleaved.
● The cleavage furrow does not penetrate the yolky portion of the cytoplasm because the yolk
platelets impede membrane formation there.
● Insect eggs have yolk in the center i.e., they are centrolecithal, and the divisions of the
cytoplasm occur only in the rim of cytoplasm, around the periphery of the cell i.e., superficial
cleavage.
● The eggs of birds and fish have only one small area of the egg that is free of yolk telolecithal
eggs, and therefore the cell divisions occur only in this small disc of cytoplasm, giving rise to
discoidal cleavage.
● Yolk is just one factor influencing a species’ pattern of cleavage.
● The importance of this inheritance can readily be seen in isolecithal eggs.
● In the absence of a large concentration of yolk, holoblastic cleavage takes place.
Four major patterns of this cleavage type can be described: radial, spiral, bilateral, and rotational
holoblastic cleavage.
Gastrulation:
● During gastrulation, the cells of the blastula are given new positions and new neighbors and
the multilayered body plan of the organism is established.
● The cells that will form the endodermal and mesodermal organs are brought to the inside of
the embryo, while the cells that will form the skin and nervous system are spread over its
outside surface.
● Thus, the three germ layers—outer ectoderm, inner endoderm, and interstitial mesoderm—are
first produced during gastrulation.
● In addition, the stage is set for the interactions of these newly positioned tissues.
● Gastrulation usually proceeds by some combination of several types of movements.
● These movements involve the entire embryo, and cell migrations in one part of the
gastrulating embryo must be intimately coordinated with other movements that are taking
place simultaneously.
● Although patterns of gastrulation vary enormously throughout the animal kingdom, all of the
patterns are different combinations of the five basic types of cell movements—invagination,
involution, ingression, delamination, and epiboly.
● In addition to establishing which cells will be in which germ layer, embryos must develop
three crucial axes that are the foundation of the body: the anterior-posterior axis, the
dorsal-ventral axis, and the right-left axis.
● The anterior-posterior AP or anteroposterior axis is the line extending from head to tail or
mouth to anus in those organisms that lack a head and tail.
● The dorsal-ventral DV or dorsoventral axis is the line extending from the back (dorsum) to the
belly (ventrum).
● The right-left axis separates the two lateral sides of the body.
● Although humans for example may look symmetrical, recall that in most of us, the heart is in
the left half of the body, while the liver is on the right. Somehow, the embryo knows that
some organs belong on one side and other organs go on the other
● Morula, a solid mass of blastomeres resulting from a number of cleavages of a zygote, or
fertilized egg. Its name derives from its resemblance to a mulberry Latin: morum.
● A morula is usually produced in those species the eggs of which contain little yolk and,
consequently, undergo complete cleavage.
● Blastula, the hollow sphere of cells, or blastomeres, is produced during the development of an
embryo by repeated cleavage of a fertilized egg.
● The cells of the blastula form an epithelial covering layer, called the blastoderm, enclosing a
fluid-filled cavity, the blastocoel.
● A single-celled zygote thus will undergo multiple rounds of cleavage, or cell division, in order
to produce a ball of cells, called a blastula, with a fluid-filled cavity in its center, called a
blastocoel.
The blastula eventually organizes itself into two layers:
● The inner cell mass which will become the embryo and
● the outer layer or trophoblast which will become the placenta; the structure is now called a
blastocyst.
Studying the chick embryo
● Pander discovered that the embryo was organized into germ layers
○ three distinct regions of the embryo that give rise through epigenesis to the
differentiated cell types and specific organ systems.
● These three layers are found in the embryos of most animal phyla:
○ The ectoderm generates the outer layer of the embryo. It produces the surface layer
epidermis of the skin and forms the brain and nervous system.
○ The endoderm becomes the innermost layer of the embryo and produces the
epithelium of the digestive tube and its associated organs including the lungs.
○ The mesoderm becomes sandwiched between the ectoderm and endoderm. It
generates the blood, heart, kidney, gonads, bones, muscles, and connective tissues.
● Pander also demonstrated that the germ layers did not form their respective organs
autonomously.
● Rather, each germ layer “is not yet independent enough to indicate what it truly is; it still
needs the help of its sister travelers, and therefore, although already designated for different
ends, all three influence each other collectively until each has reached an appropriate level.”
● Pander had discovered the tissue interactions that we now call induction.
● No vertebrate tissue is able to construct organs by itself; it must interact with other tissues.
● Meanwhile, Rathke followed the intricate development of the vertebrate skull, excretory
systems, and respiratory systems, showing that these became increasingly complex.
● He also showed that their complexity took on different trajectories in different classes of
vertebrates.
● For instance, Rathke was the first to identify the pharyngeal arches.
● He showed that these same embryonic structures became gill supports in fish and the jaws
and ears among other things in mammals