Developmental Biology

(ZOO-409)
Credit Hours: 3

Developmental Patterns
Lecture # 3
• After fertilization, the development of a multicellular organism
proceeds by a process called cleavage, a series of mitotic
divisions whereby the enormous volume of egg cytoplasm is
divided into numerous smaller, nucleated cells. These
cleavage-stage cells are called blastomeres.
• In most species (mammals being the chief exception), the rate of cell division and
the placement of the blastomeres with respect to one another is completely under
the control of the proteins and mRNAs stored in the oocyte by the mother.
• During cleavage, however, cytoplasmic volume does not increase. Rather, the
enormous volume of zygote cytoplasm is divided into increasingly smaller cells.
• First the egg is divided in half, then quarters, then eighths, and so forth. This
division of egg cytoplasm without increasing its volume is accomplished by
abolishing the growth period
between cell divisions (that is, the G and G phases of the cell
1 2

cycle).
• Meanwhile, the cleavage of nuclei occurs at a rapid rate never seen again (not even
in tumor cells). A frog egg, for example, can divide into 37,000 cells in just 43
hours.
• One consequence of rapid cell division is that the ratio of
cytoplasmic to nuclear volume gets increasingly smaller as
cleavage progresses. In many types of embryos, this decrease in
the cytoplasmic to nuclear volume ratio is crucial in timing the
activation of certain genes.
• For example, in the frog, transcription of new messages is not
activated until after 12 divisions. At that time, the rate of cleavage
decreases, the blastomeres become motile, and nuclear genes
begin to be transcribed. This stage is called the mid-blastula
transition.
• Cleavage begins soon after fertilization and ends shortly after
the stage when the embryo achieves a new balance between
nucleus and cytoplasm.
• The transition from fertilization to cleavage is caused by the
activation of mitosis promoting factor (MPF).
• Blastomeres generally progress through a cell cycle consisting
of just two steps: M (mitosis) and S (DNA synthesis)
• Gerhart and co-workers (1984) showed that MPF undergoes
cyclical changes in its level of activity in mitotic cells. The MPF
activity of early blastomeres is highest during M and
undetectable during S. Newport and Kirschner
(1984) demonstrated that DNA replication (S) and mitosis (M)
are driven solely by the gain and loss of MPF activity.
• Cleavage is actually the result of two coordinated processes.
The first of these cyclic processes is karyokinesis—the mitotic
division of the nucleus. The mechanical agent of this division is
the mitotic spindle, with its microtubules composed
of tubulin (the same type of protein that makes up the sperm
flagellum).
• The second process is cytokinesis—the division of the cell. The
mechanical agent of cytokinesis is a contractile
ring of microfilaments made of actin
Patterns of embryonic cleavage
• Different organisms undergo cleavage in distinctly different
ways. The pattern of embryonic cleavage particular to a species
is determined by two major parameters:
the amount and distribution of yolk protein within the cytoplasm,
and factors in the egg cytoplasm that influence the angle of the
mitotic spindle and the timing of its formation.
• The amount and distribution of yolk determines where cleavage
can occur and the relative size of the blastomeres. When one
pole of the egg is relatively yolk-free, the 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
• Pattern of embryonic cleavage is determined both by the position of the
mitotic spindles and by the amount and distribution of yolk.
• Yolk tends to inhibit cleavage. It slows down or actually prevents complete
cleavage.
• Yolk is an adaptation of those animals that go through more or less of
embryogenesis isolated from any food supply.
• Some animals, like sea urchin, have relatively little yolk because they
rapidly develop into a free swimming larval form that acquires nutrients
from their environment. Other animals such as marsupials are born
prematurely, but are provided nourishment in a parental pouch.
• Placental mammals develop a specialized organ through which the embryo
is nourished throughout development and so also have little yolk.
• The types of eggs based on yolk characteristics are described as:
Isolecithal: sparse evenly distributed yolk, eg., sea urchin, mouse
Mesolecthal: moderate amount of yolk, often unevenly distributed,
eg., frog
Telolecithal: dense yolk concentrated at one end, eg., bird, reptile
Centrolecithal: yolk concentrated at the middle of the egg, eg. fly
• Many eggs are polarized with a yolk rich pole, termed the vegetal
pole and a yolk poor pole termed the animal pole, e.g., frog.
• The zygotic nucleus is generally displaced towards the animal pole.
Zygotes with relatively little yolk (isolecithal and mesolecithal)
cleave holoblastically. The cleavage furrow extends all the way
through the egg.
• While telolecithal and centrolecithal zygotes
undergo meroblastic cleavage where the cleavage plane extends
only to the accumulated yolk. In centrolecithal eggs (many insect
eggs) cleavage is meroblastic and superficial, while in telolecithal
eggs (birds and fish) cleavage is discoidal.
Holoblastic cleavage
• The holoblastic type of cleavage is commonly seen in eggs
containing moderate to sparse amount of yolk. Examples
of animals with eggs that divide holoblastically include the
amphibians, mammals, echinoderms,
annelids, flatworms, nematodes, etc.
Meroblastic cleavage
• Meroblastic cleavage takes place in those eggs where the
amount of yolk is high, which leads to the partial or incomplete
cleavage. It is basically of two types
• Spherical
• discoidal.
• The discoidal type of cleavage takes place in the reptiles and birds
whose eggs are telolecithal in nature. The cleavage furrow does
penetrate the yolk. In superficial cleavage, the cytokinesis does not
take place and it leads to a polynuclear egg
• There are several types of cleavage symmetry seen in nature:
radial(echinoderms, amphibians), spiral (mollusks, annelids),
Bilateral (ascidians,tunicates), Rotational (mammals).
• It is suggested that there is some factor in the vegetal pole of
the egg that determines the formation of the micromeres and
further that there must be a “molecular clock” that starts at egg
activation. The clock is independent of the actual cleavage
event.
• Two theories attempt to account for the pattern of enlargement of the
blastocyst

1. The osmotic theory suggests that ions and proteins are secreted into the
blastocoel by the blastomeres and this results in a pressure buildup due to the
osmotic flow of water. This pressure would then be responsible for aligning the
axis mitosis of the blastomeres and the enlargement of the blastocoel.

2. The alternate theory by Wolpert and his colleagues suggests that it is really
the adhesive interactions among the blastomeres and between the blastomeres
and the hyaline layer that aligns the mitotic axis's. That is the adhesion to the
hyaline is greatest, the adhesion to other blastomeres is next, and finally the
interaction with the blastocoel wall is least. The dominant adhesion with the
hyaline layer forces the expansion of the blastocyst and blastocoel.
• The cells of the blastula grow cilia on their outer surface,
secrete a “hatching enzyme” (hyalinase) and become free
swimming.
Gastrulation
• Gastrulation is the process of highly coordinated cell and tissue
movements whereby the cells of the blastula are dramatically rearranged.
• The blastula consists of numerous cells, the positions of which were
established during cleavage.
• During gastrulation, these cells 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 inside 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.
• The movements of gastrulation involve the entire embryo, and
cell migrations in one part of the gastrulating embryo must be
intimately coordinated with other movements occurring
simultaneously. Although the patterns of gastrulation vary
enormously throughout the animal kingdom, there are only a
few basic types of cell movements. Gastrulation usually
involves some combination of the following types of
movements.
• Invagination. The infolding of a region of cells, much like the indenting of a
soft rubber ball when it is poked.
• Involution. The inturning or inward movement of an expanding outer layer
so that it spreads over the internal surface of the remaining external cells.
• Ingression. The migration of individual cells from the surface layer into the
interior of the embryo.
• Delamination. The splitting of one cellular sheet into two more or less
parallel sheets.
• Epiboly. The movement of epithelial sheets (usually of ectodermal cells)
that spread as a unit, rather than individually, to enclose the deeper layers
of the embryo
 LAW OF RECAPITULATION
• The "law of recapitulation," also known as the biogenetic law or
embryological parallelism, is a concept that was proposed by German
biologist Ernst Haeckel in the 19th century. Haeckel suggested that the
development of an individual organism (ontogeny) recapitulates or mirrors
the evolutionary development of its species (phylogeny).

• In other words, according to the law of recapitulation, the stages of