Biology 81: Integrated Biology

Animal Development
Phases of Embryonic Development
Fertilization
Fertilization
Fertilization occurs when the sperm and
the egg fuse to form the diploid zygote.

For mammals and many terrestrials, the

fertilization process involves:

1. The Acrosomal Reaction.

2. The Egg Depolarization/The

Cortical Reaction.
 A Detail on Sperm Structure
The sperm cell has a head which contains the nucleus, the acrosome, and
the centriole.
 A Detail on Sperm Structure
The middle piece contains the mitochondria, and the tail is mostly
plasma membrane.
 A Detail on Egg Structure
The egg cell is surrounded by the corona radiata (a layer of cells that
gives the egg nutrients) and the zona pellucida (a jelly-like layer that
coats the egg).
 A Detail on Egg Structure
Within the egg cell is the nucleus (which still hasn’t finished meiosis II),
the cortical granules (vesicles found on the inner side of the cell
membrane), and the cytoplasm.
 The Acrosomal Reaction
First, the acrosomal reaction!

Green is the acrosome

Blue is the nucleus

Red is mitochondria
 The Acrosomal Reaction
As the sperm gets near the egg, it has to swim through the corona radiata
cells. The friction of the cells causes the acrosome head of the sperm to
wear down.
 The Acrosomal Reaction
When the sperm gets near the egg, the acrosome of the sperm binds to
the zona pellucida of the egg.

The acrosome head breaks and releases enzymes that can “eat” through
the zona pellucida, allowing the nucleus of the sperm to get to the egg.
 The Acrosomal Reaction
After the nucleus of the sperm gets to the egg, actin microfilaments
protrude out and bind to receptors on the egg’s membrane.

The membranes of the egg and sperm fuse together, which allows for the
entry of the sperm nucleus.
 Preventing Polyspermy
But lots of sperm usually get to one egg. How do you ensure that only
ONE sperm nucleus fertilizes the egg?

(Why would it be bad to have

multiple sperm fertilize the egg?)
 Depolarization
Binding of the sperm to the egg receptors triggers opening of Na+
channels.

Since the concentration of Na+ is higher outside the cell, an influx of

Na+ triggers the zygote to depolarize.

This is a FAST reaction to prevent polyspermy (msecs).

 Cortical Reaction
The receptors that the sperm binds to include RTKs!

The binding triggers the IP3 pathway to open Ca+2 ion channels that
cause calcium to exit the ER.
 Cortical Reaction
The release of calcium into the cytosol triggers the cortical granules to
fuse with the zygote’s plasma membrane and release their contents.
 The Cortical Reaction
The released contents of the cortical granules form the fertilization
envelope, a protein coat that expands away from the zygote surface to lift
non-fertilizing sperm away. Other sperm cannot got through this
envelope.

The fertilization envelope acts as the slow block to polyspermy (takes

~10 seconds to fully develop).
 The Calcium Reaction
The reaction occurs first at the acrosomal point and then spreads out
over the whole zygote.
Creating the Fertilization Envelope
Fertilization
After fertilization, the ovum
completes meiosis II, and the
resulting cell is known as a zygote.
The Zygote: A Fertilized Egg
A fertilized egg is known as a zygote and is diploid.
(in humans, 46 chromosomes)
(haploid egg + haploid sperm = diploid zygote)
Notes on Fertilization
Overall, the process of fertilization is
relatively slow in mammals; the first
cell division occurs 2–36 hours after
sperm binding in mammals.
Triggering Embryogenesis
The rise in Ca2+ in the cytosol of the egg
also increases the rates of cellular
respiration and protein synthesis by the
egg cell.

With these rapid changes in metabolism,

the egg is said to be activated.

The sperm nucleus merges with the egg

nucleus and cell division begins.
Cleavage
Cleavage
The zygote undergoes rapid cell
division without any cell growth.
This is known as cleavage.

Thus, after 3 cleavages, at the 8 cell

stage, and the resulting embryo is
the same size as the original zygote.

Remember the need for polar bodies?

 Importance of Polar Bodies

So you need a large zygote to

start off with for all of the
rapid divisions to occur.
Blastula Formation
Blastula Formation
After 7 cleavages, the resulting
embryo of 128 cells is known as the
blastula.

Note that the blastula is still the same

size as the zygote.
Cleavage and Embryo Size
Blastula Formation
The way the blastula forms varies
between organisms.
 Blastula Formation With Yolks
In frogs and many other land animals, cleavage is asymmetric due to the
distribution of yolk (stored nutrients).

The vegetal pole has more yolk; the animal pole has less yolk.

The yolk greatly affects the pattern of cleavage.

Blastula Formation With Yolks
The first two cleavage furrows
in the frog form four equally
sized blastomeres.

The third cleavage is

asymmetric, forming unequally
sized blastomeres.

This asymmetry is due to the

yolk in the vegetal hemisphere.
Division of a Fertilized Yolky Egg
Blastula in Humans
Human eggs have very little yolk.

A blastocyst is the human equivalent of

the blastula.
Blastula in Humans
The inner cell mass is a cluster of cells
at one end of the blastocyst (these are
embryonic stem cells).

The trophoblast is the outer epithelial

layer of the blastocyst and does not
contribute to the embryo, but instead
initiates implantation.
Blastula in Humans
It is during blastula formation that cells
lose their totipotency, or the ability to
turn into any type of cell or a complete
embryo.
Blastula in Humans
It is during blastula formation that the
zygote normally reaches the uterus for
implantation.

(Note: it still takes ~a week after

implantation before your body has
enough hormone change to test positive
on a pregnancy test).
Development in Humans
Development in Humans
 Extra Protection For the Zygote
Following implantation, the trophoblast continues to expand, and a set of
extraembryonic membranes is formed.

These enclose specialized structures outside of the embryo and act as life
support.

Land vertebrates form four extraembryonic membranes: the chorion,

allantois, amnion, and yolk sac.
Membrane Roles

- Gas exchange

- Gets rid of waste

- Prevents drying out

- Encloses yolk
(if present)
 Egg Shells
Specifically for organisms that have external zygote development, the
extraembryonic membranes could form into eggs for birds and other
reptiles.
Morphogenesis
 Morphogenesis
After cleavage, the rate of cell division slows down and morphogenesis
begins.

Morphogenesis, the process by which cells occupy their appropriate

locations, involves:

Gastrulation, the movement of cells from the blastula surface to the

interior of the embryo.

Organogenesis, the formation of organs.

Gastrulation
In gastrulation, cells migrate inward
to form an archenteron, a pocket that
will eventually become the digestive
tract.

The opening hole of the archenteron

is known as the blastopore, and
eventually becomes the anus of most
organisms.
 Archenteron and Germ Layers
The formation of the archenteron designates cells to form germ layers:
the endoderm, mesoderm, and ectoderm.

ectoderm

mesoderm endoderm
In Humans, the 3 Germ Layers Become:
 Organogenesis
During organogenesis, various regions of the germ layers develop into
rudimentary organs.

Adoption of particular developmental fates is called determination, and

may cause cells to change shape or even migrate to a new location in
the body.
 Organogenesis
Organogenesis starts with neurulation. The “top” part of the zygote
mesoderm forms into a notochord, which is a rod that goes down the
whole dorsal side of the embryo.
 Neurulation
The notochord secretes signaling molecules (like Sonic the Hedgehog
aka SHH) to the area above it, causing the ectoderm above to form the
neural tube.

This is an example of induction, when cells or tissues cause a

developmental change in nearby cells.
 The Neural Tube
The neural tube will become the central nervous system (brain and spinal
cord).

The notochord disappears before birth, but contributes to parts of the

discs between vertebrae.
 Cell Migration
After neural tube formation, neural crest cells develop along the neural
tube of vertebrates and migrate in the body, eventually forming various
parts of the embryo (nerves, parts of teeth, skull bones, and so on).

Mesoderm lateral to the notochord forms blocks called somites.

Parts of the somites dissociate to form mesenchyme cells, which migrate

individually to new locations.
 Notes on Morphogenesis
Morphogenesis in animals involves movement of cells.

In animals, movements of parts of a cell can bring about cell shape

changes, or can enable a cell to migrate to a new location.

The microtubules and microfilaments of the cytoskeleton are essential to

these events.

Apoptosis and reorganization of the cytoskeleton is a major force in

changing cell shape during development.
Videos of Development (The Frog)
Videos of Development (Chicken)
 Similarities in Development
Some species look very similar in
their early embryonic stages.
Additionally, their embryonic cells
develop in the same order with
similar patterns to produce
homologous structures.

For example, on the right, these

vertebrates all begin looking like a
guppy, and develop their limbs/fins,
spine, organs, and brains in a
similar fashion.
 Model Organisms
Biologists use model organisms to study development, chosen for the
ease with which they can be studied in the laboratory.

Development occurs at many points in the lifecycle of an animal.

Across a range of animal species, embryonic development involves

common stages that occur in a set order.
Similarities in Development

Pharyngeal
pouches

Post-anal
tail

Chick embryo Human embryo

Other Notes
 Determination vs Differentiation
Determination is the term used to describe the process by which a cell or
group of cells becomes committed to a particular fate.

(These stem cells will BECOME the eye)

Differentiation refers to the resulting specialization in structure and

function.

(These spermatocytes will grow a tail to mature into sperm cells)

 Genes and Development
Cells in a multicellular organism share the same genome.

Differences in cell types are the result of the expression of different sets
of genes.
 Fate Mapping
Fate maps are diagrams showing organs and other structures that arise
from each region of an embryo.

Classic studies using frogs indicated that cell lineage in germ layers is
traceable to blastula cells.
 Studies with C. elegans
Later studies of C. elegans used the ablation (destruction) of single cells
to determine the structures that normally arise from each cell.

The researchers were able to determine the lineage of each of the 959
somatic cells in the worm.
Studies with C. elegans
 Axis Formation
A body plan with bilateral symmetry is found across a range of animals.

This body plan exhibits asymmetry across the dorsal-ventral and

anterior-posterior axes.

The right-left axis is largely symmetrical.

 Axis Formation in Birds vs Mammals
In chicks, gravity is involved in establishing the anterior-posterior axis.

Later, pH differences between the two sides of the blastoderm establish

the dorsal-ventral axis.

In mammals, experiments suggest that orientation of the egg and sperm

nuclei before fusion may help establish embryonic axes.

In insects, morphogen gradients establish the anterior-posterior and

dorsal-ventral axes.
Questions?
 The “Experiment”
Can baby birds develop normally in space?
Gravity in Space?
 Attendance
Which receptors found on the egg surface triggers the cortical reaction
when bound?

A. GPCRs
B. Ligand-gated ion channels
C. RTKs
D. Neuroreceptors