INTRODUCTION TO DEVELOPMENTAL BIOLOGY

First research into embryology

• Aristotle recognised the 3 types of reproductive strategy

o Cracked open chicken eggs every day during a 3-week incubation period and examined the embryo
using only the naked eye

3 types of reproductive strategy:

• OVIPARITY
o Eggs are laid
o Little to no embryonic development in the mother
o Internal/ external fertilisation
o E.g. invertebrates, fish, amphibians, reptiles, birds and monotreme mammals

• OVOVIVIPARITY:
o Eggs retained in the mothers body until ready to hatch.
o No placental connection to the mother
o Internal fertilization
o Young are born live
o E.g. invertebrates, fish, amphibians and reptiles.

• VIVIPARITY:
o Eggs retained in the mother’s body until ready to hatch
o Placental connection to mother
o Internal fertilization
o Young are born live
o E.g. Most mammals
▪ Marsupial mammals are born in a very immature state and lack a complex placenta

2 theories of how different parts of the embryo are formed?

• PREFORMATION: organs and tissues are preformed and correctly positioned in the fertilized egg. Organs and
tissues enlarge during embryonic development.
o Homunculus (Little Man): Preformationists believed that a small, fully-formed human was visible in
the sperm head
o If true, male sperm would require sperm of their own, and the mother would make no contribution
to the next generation
o Disputed by the fact that children can inherit their mother’s characteristics

• EPIGENESIS: organs and tissues are formed gradually and the complexity of the embryo increases with time.
o Theory was supported with the development of better microscopes and staining techniques showing
increasing complexity in structures with time
▪ Led to discovery of intermediate stages that varied significantly
▪ MORPHOGENESIS: Movement of cells resulting in tissue formation
▪ Embryonic stem cells undergo morphogenic movement

Sequences of development

• Comparative embryologists identifies that embryos from different species pass through similar stages of
development
o Fertilisation followed by cleavage divisions
o Formation of a fluid-filled sphere called a blastula
▪ Blastula is where embryonic stem cells come from
• Cells are capable of giving rise to all tissue types
What can cells differentiate into?

• TOTIPOTENT: Cell can give rise to all embryonic and extra-embryonic tissues
• Cells can undergo lineage restriction to become pluripotent
• PLURIPOTENT: Cell can give rise to only embryonic tissues
• Cells undergo a second lineage restriction event called gastrulation in which the blastula forms a gastrula
• MULTIPOTENT: Ectoderm can no longer give rise to all embryonic tissue
o The gastrula differentiates into the ectoderm, mesoderm, endoderm or germ cells
o Gastrula can be pattered to develop a head/tail, dorsal/ventral, medial/lateral
• Cells undergo a final lineage restriction event called differentiation in which multipotent stem cells form
progenitor cells that have a tendency to differentiate into a specific type of cell
o Progenitor cells can only divide a limited number of times

▪ During gastrulation, stem cells undergo morphogenic movement to generate 3 germ layers
o Development of multi-layered
larva by gastrulation
o 3 germ layers become observable
(ectoderm, mesoderm, endoderm)
▪ Layers present in all
triploblastic embryos
▪ Some phyla (e.g. porifera,
cnidarians, and
ctenophores) are
diploblastic and lack true mesoderm
o Through improved microscopes and staining techniques, histologists can recognise over 200 cell
types in humans formed by differentiation

3 germ layers

Phylotypic stage

• Latest stage in development when different species are not distinguishable

• Important because evolution works by adjusting developmental biology

Vertebrate body plan

• Similar arrangements of tissues and organs can be found in all vertebrate embryos at the phylotypic stage
o Only amniotic membrane and umbilical cord are not found in all vertebrates
 o f = forebrain
m = midbrain
h = hindbrain
t = trunk
h = heart
uc = umbilical cord
pa = pharyngeal arches
am = amniotic membrane
e = epidermis
nt = neural tube
n = notochord
i = intermediate mesoderm
a = aorta
lp = lateral plate
ua = umbilical artery
g = gut epithelium.

Pharyngeal arches

• Also known as the branchial, visceral and gill arches

• Found at phylotypic stage of all vertebrate embryos
• Become gills in fish, and jaws, ears and neck in mammals in a lineage
restriction event

How do cells acquire their different characteristics?

• Acquired through a process called mosaic development

o August Weismann (1834-1914)
o Proposition that eggs contain a mosaic of different determinants that direct the future development
of cells
▪ Examples of determinants include those for the skin, nervous system, muscle, blood etc.
▪ Preformation at the molecular level
o Factors are localized to different parts of the egg cytoplasm
o Asymmetric distribution of factors occurs during cleavage

Yellow cytoplasm experiment

• Edwin Conklin (1863-1952)

• Principle organs of the larva in their positions and proportions can be
marked out in the 2-cell stage by distinct kinds of protoplasm
• Experimentation with ascidian eggs showed that some species had
different coloured cytoplasms
o Fate map was produced by following development of the
cytoplasms
o E.g. yellow cytoplasm was always found in muscle cells of the
larva, suggesting that it contained a muscle determinant

Birth of Experimental Embryology

• Wilhelm Roux (1850-1924)

• Experimentation with frog embryos
• Single cell of a frog embryo in its 2-cell phase was killed using a hot needle
• Remaining cell formed only half an embryo
o Led to the conclusion that the frog egg was a mosaic of different self-differentiating parts as
remaining cell received only a subset of the egg’s determinants
Regulative Development

• Hans Driesch (1867-1941)

• Competes with mosaic theory
• Separation of cells in the 4-cell phase of an embryo results in the
formation of 4 small, complete larvae

• Hans Spemann (1869-1941)

• Use of a fine hair as a ligature to separate 2-cell newt embryos
into individual cells
• Each cell formed a half-sized larva

How were these contradictions overcome?

• Standard dichotomy between the two experiments

o In the first experiment, the death of 1 out of 2 cells led to the development of only half an
embryo
o In the second experiment, the separation of 4 cells led to the development of 4 complete
embryos
• When the cell is still in contact with the other cell (proteins in contact with membrane) therefore the
other cell is unaware that the cell is dead
• When cells are separated, there is no longer contact between cells, therefore the cell is aware that it
must develop into the full embryo
• This suggests that cells communicate with each other when deciding what each cell will form
Amphibian Organizer

• Hans Spemann (1869-1941) & Hilde Mangold (1898-

1924)
• Experiment involving the transplant of the notochord
(cells that develop into the nervous system and head)
of a newt gastrula into the opposite side of a similarly
staged host
o Results in development of 2 nervous systems
and 2 heads
o Newt species with different pigmentations
used to track the transplanted cells
• Resulted in the for mation of a conjoined twin
o Transplanted cells formed the notochord and
partially the nervous system
o Host ectodermal cells formed remaining
nervous system instead of forming the
epidermis
• Transplanted cells changed the fate of host cells
• Process called neural induction

Neural Induction

• During gastrulation, dorsal mesoderm (notochord) moves beneath the dorsal ectoderm and released signals
to induce the nervous system
o Absence of these signals results in the dorsal ectoderm forming the epidermis
o In ‘amphibian organizer’ experiment, the notochord was introduced on the ventral side of the
embryo and lead to formation of a second nervous system

Embryonic Induction

• INDUCTION: the process whereby a cell or tissue signals to another cell or tissue in the embryo thus affecting
the development of the responding cell or tissue.
• COMPETENCE: the ability of a cell or tissue to respond to an inducing signal
o Embryonic tissues only remain competent for a limited period of time.

Cell Fate

• Cell fate is determined over several different steps that restrict potency
o POTENCY: Range of cell types a cell can form
o E.g. In neuron formation, cell fate is determined by forming ectoderm (instead of mesoderm/
endoderm), neuroepithelium instead of epidermis, and form a neuron instead of a support cell
• Early decisions likely involve cytoplasmic determinants whereas later decisions involve cell-cell interactions
(induction)