Chapter 47 Presentation

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Chapter 47
Animal Development
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The Zygote
The question of how a
zygote becomes an
animal has been
asked for centuries.
 As of the 18th century,
preformation was the
prevailing notion. The
idea was that a
preformed miniature
infant (homuniculus)
was contained within
an egg or a sperm.

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The Zygote
More than 2000 years ago: Aristotle
originally posed the idea of epigenesis
to explain embryonic development.
 This idea explained that an animal
gained form gradually from a relatively
formless egg.

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The Zygote
As microscopy improved in the 19th
century, embryologists began to see
that embryonic development took place
in a series of progressive steps and
epigenesis became the favored
explanation.
 The genome of the zygote and
differences in early embryonic cells
determine an organism’s development.

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The Zygote
There are some important events that
regulate animal development:
 1. Fertilization
 2. Cleavage
 3. Gastrulation
 4. Organogenesis

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1. Fertilization
Fertilization is when an egg and a
sperm unite forming a zygote.
 The main function of fertilization is to
produce a diploid egg.
 Egg activation occurs when the sperm
contacts the egg’s surface and initiates
metabolic reactions that trigger the
onset of embryonic development.

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1. Fertilization

In general, when eggs meet sperm, an
acrosomal reaction is triggered.
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1. Fertilization

The process begins when a vesicle at
the tip of the sperm called the acrosome
discharges hydrolytic enzymes.
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1. Fertilization

These enzymes dissolve the egg’s jelly coat
enabling the acrosomal process to penetrate
the it and attach to the egg’s surface.
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1. Fertilization

The sperm binding receptors on the
egg’s surface are attached to the
vitelline layer.
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1. Fertilization

This recognition of the egg’s surface receptors to
sperm is what ensures only sperm from the
same species can penetrate the egg.
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1. Fertilization

The binding of the sperm membrane to
the membrane of the egg changes the
state of the ion channels of the egg.
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Fast Block to Polyspermy

Within 1-3 seconds, depolarization occurs where Na+
ions rush into the egg, and is known as the fast block
to polyspermy. It prevents multiple sperm from
entering the egg.
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Fast Block to Polyspermy

The fast block to polyspermy is shortlived and the membrane polarization
only lasts about one minute.
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Slow Block to Polyspermy

The fusion of the sperm and egg also triggers
a series of changes in the egg that are more
long-lived.
 Sperm binding activates a signal transduction
pathway which causes calcium ions to be
released from the egg’s ER into the cytosol.
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Slow Block to Polyspermy

Release of Ca2+ from the ER occurs 1st
at the site of sperm entry and moves
like a wave over the surface of the egg.
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Slow Block to Polyspermy

The high concentration of calcium initiates the
cortical reaction.
 The reaction triggers the fusion of the egg’s
plasma membrane with numerous vesicles
lying just beneath the membrane.
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Slow Block to Polyspermy
The contents of these vesicles enter the
previtelline space.
 Enzymes in these contents degrade the
proteins holding the vitelline layer to the
plasma membrane.

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Slow Block to Polyspermy
These changes transform the vitelline
layer into the fertilization envelope.
 No more sperm can enter the egg at
this time.

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Slow Block to Polyspermy
This is referred to as the slow block to
polyspermy.
 The sharp rise in Ca2+ also increases
the metabolism of the egg.

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Slow Block to Polyspermy
After about 20 minutes, the egg nucleus
fuses with the sperm nucleus creating a
diploid zygote.
 Fertilization has many common features
among species and many differences.
These differences are mainly with timing
and various stages of meiosis.

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1. Fertilization--Sperm Activation

In mammals, secretions in the female
reproductive tract to help to activate sperm.
 Additionally, the mammalian egg is cloaked
by follicle cells through which the sperm must
penetrate before reaching the zona pellucida.
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1. Fertilization--Sperm Activation

The zona pellucida is the
external matrix of the egg
and functions as a sperm
receptor.
 When sperm binds to
this, it induces many of
the same reactions as
seen in the sea urchin.
 One difference is that
there is no fast block to
polyspermy in mammals.
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1. Fertilization--Sperm Activation
After the egg and sperm membranes
fuse the whole sperm is taken into the
egg.
 The egg lacks a centrosome which the
sperm has.
 The centrosome will duplicate and
assist in the creation of the mitotic
spindle which will be used for the first
cell division.

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1. Fertilization--Sperm Activation
In mammals, the nuclei do not
immediately fuse. Instead, the 2 sets of
c-somes share a common spindle
apparatus during the 1st mitotic division.
 Only after the first division, as diploid
nuclei form in the 2 daughter cells, do
the chromosomes from the 2 parents
come together in a common nucleus.

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1. Fertilization
Fertilization is quite slow in mammals.
 Once fertilization is complete, rapid cell
division ensues and the cells proceed
through the M and S phases of the cell
cycle virtually skipping G1 and G2.

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1. Fertilization

Thus, the embryo doesn’t enlarge
much. Rather, it becomes an aggregate
of blastomeres called a blastocyst
(blastula).
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Copyright ©2005 Pearson Education, Inc. Publishing as Pearson Benjamin Cummings. All rights reserved.
2. Cleavage

The color of the frog
egg cells allows for
easy following of cells
during development.
 Animal hemisphere is
deep gray due to
melanin.
 The vegetal pole is
yellow due to lack of
melanin.
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2. Cleavage

The first 5 to 7 divisions produce what is
called the morula.
 Within the morula a fluid filled cavity called a
blastocoel forms. When fully formed, the
aggregate of cells is now called blastula.
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2. Cleavage

Many animals, with the exception of
possibly mammals, have zygotes with a
definite polarity that determines the
pattern of cleavage.
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2. Cleavage

In classic examples
such as the frog, the
distribution of yolk in the
egg determines
cleavage patterns.
 The vegetal pole
contains a large amount
of yolk, and cleavage
through this is slow.
 The animal pole
contains very little yolk
and cleavage occurs
quickly through it.
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2. Cleavage--3 Body Axes
The 3 body axes are
established early on
in development.
 1. Dorsal/ventral
 2. Right/left
 3. Anterior/posterior

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2. Cleavage
Using the frog as an
example, fusion of
the sperm and egg
initiates a body axis.
 At this point, the
animal pole will
move toward the
vegetal pole passing
through the site of
sperm entry.

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2. Cleavage

As a result of the
cortical rotation, a
light gray region
known as the gray
crescent forms in
some species and
the dorsal/ventral
axis is formed in
the zygote.
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2. Cleavage
This light gray
region forms the
dorsal side of the
embryo.
 In frogs (animals
with a thick vegetal
pole), the
blastocoel, is
located in the
animal pole
because the yolk
isn’t as thick.

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2. Cleavage

Yolk is very plentiful
in many animals
such as a bird.
 In this example, the
yolk is actually an
egg cell swollen
with yolk nutrients.
A small white disk
of actual cytoplasm
is located on the
animal pole.
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2. Cleavage

The entire cell
is surrounded
by a protein
rich solution of
egg white
which provides
additional
nutrients to the
growing
embryo.
Albumen
Chalaza
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Yolk
Disk of Cytoplasm
2. Cleavage
Cleavage is
restricted to the
small disk of
yolk free
cytoplasm.
 Cleavage can’t
proceed through
the dense yolk.

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2. Cleavage
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Incomplete division of the
yolk rich egg is known as
meroblastic cleavage.
 In contrast, holoblastic
cleavage results in
complete division of the
egg that lacks a lot of
yolk, (or a moderate
amount of yolk).

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2. Cleavage
The cleaving cells on the top of the yolk
of a bird egg produce a cap of cells
called the blastoderm.
 The blastoderm then rests on the yolk
and divides into 2 layers called the
epiblast and the hypoblast.

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2. Cleavage
The cavity between them is called the
blastocoel.
 This is the equivalent of the blastula
stage of the frog/sea urchin.

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3. Gastrulation

Gastrulation is the rearrangement of
cells of the blastula to form a 3 layered
embryo with a primitive gut.
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3. Gastrulation

Gastrulation differs between animal
groups, but it shares some general
characteristics:
 A.
Changes in cell motility.
 B. Changes in cell shape.
 C. Changes in cellular adhesion.
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3. Gastrulation

The result is that some cells
at the surface of the blastula
move inward establishing 3
cell layers.
 The 3 layered embryo is
called the gastrula.
 The 3 layers are given the
term embryonic germ layers
and consist of:
 1. Endoderm--lines digestive


tract.
2. Mesoderm--partially fills the
space between endoderm and
ectoderm.
3. Ectoderm--which forms the
outer layer of the gastrula.
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3. Gastrulation--Sea Urchin




These 3 layers will form all
cell layers and tissues in
the adult animal.
In the sea urchin, the
blastula consists of a single
layer of cells enclosing a
blastocoel.
Gastrulation begins at the
vegetal pole.
Here, migratory cells called
mesenchyme cells enter
the blastocoel.
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3. Gastrulation--Sea Urchin

The cells which remain
form the vegetal plate
which eventually buckles
inward by a process
called invagination.
 Invagination proceeds
forming the archenteron
(primitive gut).
 The open end of the
archenteron will become
the anus, it is called the
blastopore.
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3. Gastrulation--Sea Urchin
The second opening
forms as the archenteron
touches the inside of the
ectoderm on the other
side. This is the mouth
(in deuterostomes).
 A rudimentary digestive
tube is now formed.

Ectoderm
Blastocoel
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Mesenchyme Cells
Vegetal Plate
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3. Gastrulation--Frog

In the frog, gastrulation
also produces a 3
layered embryo with an
archenteron.
 Gastrulation is more
complex due to the yolk
content of the cells, and
the fact that the wall of
the blastula is more than
1 cell-layer thick.
 Gastrulation begins on
the dorsal side of the
blastula in a line along
the region of the gray
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crescent.
3. Gastrulation--Frog
This invagination
forms the dorsal lip
which pinches inward
around the cell.
 The cells on the
surface of the
embryo continue to
roll inward in a
process called
involution forming the
future endoderm and
mesoderm.

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3. Gastrulation--Frog

Eventually, the
blastocoel collapses as
the archenteron replaces
it. The archenteron is
lined with endoderm.
 As gastrulation
completes, the
blastopore encircles the
yolk plug.
 After the archenteron
reaches the opposite
side of the embryo, a
mouth will form.
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3. Gastrulation--Chick

The chick’s gastrulation is similar to that of
a frog except the inward movement of cells
during gastrulation is affected by the yolk.
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3. Gastrulation--Chick

Cleavage in the chick results in a
blastoderm with an epiblast and a
hypoblast.
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3. Gastrulation--Chick

All cells that form the embryo will come from the epiblast.
 During gastrulation, some of the epiblast cells move toward
the blastoderm’s midline producing a thickening called the
primitive streak.
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3. Gastrulation--Chick
The primitive streak will
form the bird’s
anterior/posterior axis.
 The primitive streak is
the functional equivalent
to the lip of the
blastopore in the frog,
but structures are
aligned differently.
 Inward moving cells
displace hypoblast cells
and form endoderm.

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Scott F. Gilbert Developmental Biology
3. Gastrulation--Chick

Other epiblast cells move laterally into the
blastocoel and form mesoderm.
 Epiblast cells remaining on the surface form
ectoderm.
 The hypoblast helps direct the formation of the
primitive streak and forms part of the stalk that
keeps the yolk mass connected to the embryo.
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4. Organogenesis


Organogenesis is the process by which parts of the 3
germ layers develop the rudiments of organs.
Organogenesis involves localized morphogenic
changes in tissue and cell shape versus the large
scale mass movement of cells seen in gastrulation.
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4. Organogenesis

Evidence of organogenesis is seen in the
appearance of folds splits and dense
clustering of cells.
 Easy observations are made in chordates
when the notochord, neural tube, etc. take
form.
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4. Organogenesis

The notochord is formed from dorsal
mesoderm which condenses just above
the archenteron.
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4. Organogenesis

Signals sent from the notochord to the
ectoderm cause the region of the ectoderm to
form the neural plate.
 The neural plate will curve inward forming the
neural tube which runs along the anteriorposterior axis of the embryo eventually
forming the CNS.
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4. Organogenesis
In vertebrates, the
neural crest forms
along the border
where the neural tube
pinches off from the
ectoderm.
 Neural crest cells then
migrate to various
parts of the embryo
giving rise to many
structures such as
peripheral nerves,
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teeth, skull bones,
etc.

4. Organogenesis
Lateral to the
notochord are
condensations of
mesodermal cells
arranged in the block
called somites.
 The somites give rise
to cells that migrate to
new locations forming
vertebrae and
muscles.

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Development
All vertebrates require an aqueous
environment to grow and develop.
 The evolution of structures that allow
this to happen have occurred and are
the shelled egg and uterus. These
structures protect the embryo with an
amnion which is a fluid filled sac.

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Extraembryonic Membranes
The development of mammals, birds,
and reptiles includes the formation of 4
extraembryonic membranes.
 These membranes include:

 Yolk
sac
 Chorion
 Allantois
 Amnion
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Extraembryonic Membranes

The yolk sac covers the yolk and develops
blood vessels that transplant nutrients
from the yolk to the embryo.
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Extraembryonic Membranes

The chorion is involved in gas exchange
with the surroundings.
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Extraembryonic Membranes

The allantois is a disposable sac for metabolic
wastes produced by the developing embryo.
It also functions with the chorion in respiration.
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Extraembryonic Membranes

The amnion protects the embryo in a
fluid filled cavity.
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