1. During cleavage, cells divide rapidly through mitosis but do not grow in size. Cell cycling is controlled by cyclin synthesis and degradation, which regulates the activity of MPF and progression through the cell cycle. 2. Cleavage patterns vary between species, from radial in sea urchins to spiral in snails to bilateral in tunicates. Gastrulation occurs through various cell movements and generally establishes the three body axes. 3. Cell fate specification occurs through different mechanisms in different species, including maternal cytoplasmic determinants, cell signaling, and cell-cell interactions. Axes can be established before or after fertilization.

1. An Introduction to Early Developmental
Processes
The Early Development of Sea Urchins
The Early Development of Snails
Early Development in Tunicates
Early Development of the Nematode
Caenorhabditis elegans

2. Cleavage
rapid cell division
no overall growth
no increase in volume

3. Cleavage
Cell Cycle Control
 Initially, factors stored in the egg control cleavage (the
cell cycle);
e.g. stored mRNAs
stored proteins
….. initiate the cell division
 Mitosis Promoting Factor (MPF) stimulates the cell cycle

4. Cleavage-Stage Cell Cycle -
blastomeres

5. Cleavage-Stage Cell Cycle-
blastomeres
MPF – mitosis promoting factor:
-Cyclin B – controls cdc2 activity
-cdc2 = cyclin dependent Kinase
CDK phosphorylates histones,
etc.
#Cyclin B degrades; cell division
stops
Cyclin B presence/degradation
controlled by egg cytoplasmic
proteins

6.  Mid Blastula Transition (MBT)
-the rate of cleavage decreases, the blastomeres
become motile, and nuclear genes begin to be
transcribed.
#Cleavage begins soon after fertilization and ends shortly
after the stage when the embryo achieves a new balance
between nucleus and cytoplasm.

7. Post Cleavage Cell
Cycle
Post MBT:
• Cell cycle adds two
G phases
• New mRNA
transcription
• Cell division
becomes
asynchronous

9. Cytoskeletal Mechanisms
karyokinesis & cytokinesis

10. Cleavage Patterns
Cleavage
 Rapid cell divisions
 Divisions of fertilized egg into many cells
*What influences the pattern of cleavage in a particular organism?

12. Gastrulation
 the process of highly coordinated cell and tissue movements whereby the
cells of the blastula are dramatically rearranged

14. Axis Formation

15. 1. The Early Development of Sea
Urchins
 Sea Urchin Cleavage
-holoblastic radial cleavage

17. Sea Urchin Development
-What characterizes the blastula stage?
 expanding blastocoel
 cilla develops
 embryo rotates in fertilization membrane
 formation of vegetal plate
 formation of hatching enzyme

18.  Germ Layers / Fate Maps
Germ Layers / Fate Maps
 inner = endoderm
 middle = mesoderm
 outer = ectoderm
Sea Urchin Zygote

19. Sea Urchin Cell Fate
Cells are specified by either: Asymmetric distribution of patterning molecules
into particular cells or cell-cell interactions
Mechanisms for establishing asymmetry:
1. Patterning molecules bound to egg cytoskeleton
2. Molecules actively transported along the cytoskeleton
3. Molecules become associated with one centrosome, and then follow that
centrosome into one of the two mitotic sister cells
 Once asymmetry is established, one cell can specify another (and
participate in reciprocal inductions)

20. Sea Urchin Development
At what stage are the fates of individual
cells determined?

21. Sea Urchin Development
How does gastrulation
begin?

22. Sea Urchin Development
What appears to be responsible for the ingression of primary
mesenchyme?

23. Sea Urchin Development
What appears to be responsible for the initial
invagination that occurs during gastrulation?

24. Sea Urchin Development
What happens during later stages of
invagination?

25. The Early Development of Snails
What kind of cleavage pattern characterizes these
animals?
-Spiral holoblastic

26. Snail Development
Orientation of cleavage
plane determines right or
left coiling snails

28. Snail Development
What appears to be responsible for the mosaic
development seen in molluscs?

29. Snail Development
What is the polar lobe and why is it important?

30. Snail Development
Why does removal of the D blastomere or its first
or second derivatives result in incomplete
larvae?
If D blastomeres don’t directly contribute cells to
formation of many structures why are they so
important to the formation of the same
structures?

31. Snail Development
How does gastrulation take place in snails?

32. Early Development in Tunicates
What type of cleavage
pattern do they have?
-bilateral holoblastic

34. Tunicate Development
In what way does the pigmentation in Styela partita provide
developmental information?

35. Tunicate Development
What evidence is there of autonomous
specification in tunicate blastomeres?
-transplant experiments
-RNA hybridization experiments
-altering β-catenin levels in cells

36. Tunicate Development
 What evidence is there for conditional specifiction?
-BMP signal from endoderm induces anterior cell to
become notocord-
-works through activation of Brachury gene
-FGF signal induces posterior cell to become
mesenchyme

37. Tunicate Development
When are the embryonic
axes established?
-dorsal-ventral – prior to first
cleavage
-anterior-posterior – prior to first
cleavage
-left-right – first cleavage

38. Tunicate Development
What is gastrulation like in
these organisms?

39. Early Development of the Nematode Caenorhabditis
elegans
What does C. elegans look like?

40. C. elegans
What pattern of cleavage is seen in this
nematode?

41. Axis formation
 Anterior-posterior axis formation

42. C. elegans
When does gastrulation
begin in this organism?

43. SUMMARY

44. 1. During cleavage, most cells do not grow. Rather, the volume of
the oocyte is cleaved into numerous cells. The major exceptions
to this rule are mammals.
2. The blastomere cell cycle is governed by the synthesis and
degradation of cyclin. Cyclin synthesis promotes the formation of
MPF, and MPF promotes mitosis. Degradation of cyclin
brings the cell back to the S phase. The G phases are added at
the midblastula transition.
3. "Blast" vocabulary: A blastomere is a cell derived from cleavage
in an early embryo. A blastula is an embryonic structure composed
of blastomeres. The cavity in the blastula is the blastocoel. If the
blastula lacks a blastocoel, it is a stereo blastula. A mammalian
blastula is called a blastocyst (in Chapter 11), and the invagination
where gastrulation begins is the blastopore.

45. 4. The movements of gastrulation include invagination, involution, ingression,
delamination, and epiboly.
5. Three axes are the foundations of the body: the anterior-posterior axis
(head to tail or mouth to anus), the dorsal-ventral axis (back to belly), and the
right-left axis (between the two lateral sides of the body).
6. In all four invertebrates described here, cleavage is holoblastic. In the sea
urchin, cleavage is radial; in the snail, spiral; in the tunicate, bilateral; and in
the nematode, rotational.
7. In the tunicate, snail, and nematode, gastrulation occurs when there are
relatively few cells, and the blastopore becomes the mouth. This is the
protostome mode of gastrulation.
8. Body axes in these species are established in different ways. In some,
such as the sea urchin and tunicate, the axes are established at fertilization
through determinants in the egg cytoplasm. In other species, such as the
nematode and snail, the axes are established by cell interactions later
in development.

46. 9. In the sea urchin, gastrulation occurs only after thousands of cells have formed, and
the blastopore becomes the anus. This is the deuterostome mode of gastrulation, and
is characteristic only of echinoderms and chordates.
10. In sea urchins, cell fates are determined by signaling. The micromeres constitute a
major signaling center. β-catenin is important for the inducing capacity of the
micromeres.
11. Differential cell adhesion is important in regulating sea urchin gastrulation. The
micromeres delaminate first from the vegetal plate. They form the primary
mesenchyme which becomes the skeletal rods of the pluteus larva. The vegetal plate
invaginates to form the endodermal archenteron, with a tip of secondary
mesenchyme cells. The archenteron elongates by convergent extension and is guided
to the future mouth region by the secondary mesenchyme.
12. Snails exhibit spiral cleavage and form stereoblastulae, having no blastocoels. The
direction of the spiral cleavage is regulated by a factor encoded by the mother and
placed into the oocyte. Spiral cleavage can be modified by evolution, and
adaptations of spiral cleavage have allowed some molluscs to survive in otherwise
harsh conditions.

47. 14. The tunicate fate map is identical on its right and left sides. The yellow
cytoplasm contains muscle-forming determinants; these act autonomously.
The nervous system of tunicates is formed conditionally, by interactions
between blastomeres.
15. The soil nematode Caenorhabditis elegans was chosen as a model
organism because it has a small number of cells, a small genome, is easily
bred and maintained, has a short lifespan, can be genetically manipulated,
and has a cuticle through which one can see cell movements.
16. In the early divisions of the C. elegans zygote, one daughter cell becomes
a founder cell (producing differentiated descendants) and the other becomes
a stem cell (producing other founder cells and the germ line).
17. Blastomere identity in C. elegans is regulated by both autonomous and
conditional specification.

48. Thank you!!!