Animal Development

CHAPTER 38
Chapter 38 Animal Development

 Key Concepts
 38.1 Fertilization Activates Development
 38.2 Cleavage Creates Building Blocks and Produces a Blastula
 38.3 Gastrulation Produces a Second, then a Third Germ Layer
 38.4 Gastrulation Sets the Stage for Organogenesis and Neurulation in
Chordates
 38.5 Extraembryonic Membranes Protect and Nourish the Embryo
 38.6 Development Continues throughout Life
 Fertilization Activates Development

 How do organisms develop from a single

cell?
 Early investigators assumed that a tiny
“homunculus” was encapsulated in a
human egg or sperm.
Egg and Sperm make different
contributions to the zygote

 Development begins with fertilization, the union of sperm and egg to form
a zygote.
 Egg cells are large—they contain the materials and information needed to
initiate and maintain early development.
Egg and Sperm make different
contributions to the zygote

 Many sperm are attracted to an egg, but

only one is allowed entry.
 Activation of an egg by a sperm triggers
changes in the egg cell membrane and the
release of ions.
 A series of events is triggered:
 Entry of additional sperm is blocked
 Metabolic rate rises
 Protein synthesis is initiated
 Cytoplasm is reorganized
Egg and Sperm make different
contributions to the zygote

 The sperm nucleus and centrosome are transferred to the egg.

 The centrosome plays a major role in forming the zygote centrosome and
organizing the zygote’s microtubules.
 When egg and sperm nuclei join, the fertilized egg becomes a diploid
zygote.
Polarity is Established Early in
Development – out of scope

 Anterior–posterior and dorsal–ventral polarity of an animal’s body is

established early in development by various mechanisms.

 In amphibian eggs, the yolk material is concentrated in the lower half

(vegetal hemisphere). Sperm binding sites are in the upper half (animal
hemisphere), which is pigmented near the surface.
Polarity is Established Early in
Development

 Entry of the sperm causes the cortical cytoplasm to shift, forming the gray
crescent. This sets up the polarities.

 Early experiments, in which amphibian eggs were bisected, showed that

cytoplasmic determinants in the gray crescent are necessary for
establishing polarity and development of a normal salamander.
Figure 38.1 The Gray Crescent
Cleavage Creates
Building Blocks and
Produces a Blastula
38.2
Cleavage and Blastula

 Cleavage—rapid series of cell division, but no cell growth

 Cells increase in number but get smaller and smaller.
 The cells are called blastomeres; the ball of cells is the blastula.
 Early stage events are controlled by maternal factors: cytoplasmic
determinants in the egg. mRNA from the mother, already present in the
egg, is used to direct protein synthesis.
 Cleavage and Blastula

 The cytoskeleton may

redistribute these factors; cell
divisions also distribute them to
different blastomeres.
 At some point, the fate of a
blastomere becomes restricted
or determined.
 Experiments in which
blastomeres are separated at
different stages reveal the fates
of each cell. Blastomeres can
also be dyed or labeled in
some way to follow their fates.
Cleavage and Blastula

 As cleavage continues, the zygote becomes divided into progressively

smaller blastomeres:
Cleavage and Blastula

 Complete cleavage: cytoplasm is

completely divided by each cell division
 Radial cleavage and spiral cleavage
patterns are seen in eggs with little yolk
and complete cleavage.
 The plane of cleavage is determined by
the orientation of the mitotic spindle.

 Radial cleavage at the 8-cell stage:

Cleavage and Blastula

 Spiral cleavage at the 8-cell stage:

Specific Blastomeres Generate
Specific Tissues and Organs – Out of Scope

 Blastomeres become determined—committed to specific fates—at

different times in different species.
 Mosaic development: specific blastomeres give rise to specific tissues and
organs of the adult animal
Specific Blastomeres Generate
Specific Tissues and Organs

 In many spirally cleaving eggs, a polar

lobe forms at the vegetal pole that
contains yolk and cytoplasmic
determinants.
 If the polar lobe is removed, an
abnormal larva develops.
 If the two blastomeres are
experimentally separated, two
abnormal larvae develop, each lacking
very specific structures.
Specific Blastomeres Generate
Specific Tissues and Organs

 In regulative
development,
cells can
compensate for
lost cells.
 Distribution of
cytoplasmic
determinants
happens later as
cell divisions
continue.
 This type occurs in
vertebrates.
The Amount of Yolk Affects Cleavage

 The amount of yolk in the egg impacts developmental processes.

 In moderately yolky eggs, cleavage is complete, but cells divide more slowly in
the yolky vegetal hemisphere, resulting in an animal hemisphere consisting of
more small blastomeres and a vegetal hemisphere consisting of fewer,
larger blastomeres.
The Amount of Yolk Affects Cleavage

 In very yolky eggs, the yolk does not

divide, and the embryo forms as a
small blastodisc of cleaving cells on
the yolk surface.
 In some insects such as Drosophila,
the zygote undergoes nuclear
divisions but no cytoplasm divisions.
The early embryo is a syncytium.
 As development proceeds, nuclei
migrate to the periphery of the egg,
and cell membranes form.
Cleavage in Placental Mammals is
Unique - Essential

 Placental mammals have non-yolky eggs and rotational cleavage. At the

2nd division, blastomeres divide in different planes:
Cleavage in Placental Mammals is
Unique

 In mammals, at the 4th cleavage the cells

separate into two groups:

• Inner cell mass will become the embryo

• Surrounding outer cells become a sac

called the trophoblast; at this stage, the
embryo is called a blastocyst

 Trophoblast cells secrete fluid, creating

a blastocoel. Later they contribute to
the placenta.
Cleavage in Placental Mammals is
Unique

 In mammals, fertilization occurs in

the upper oviduct; cleavage takes
place as the zygote travels down
the oviduct.
 When the blastocyst arrives in the
uterus, it adheres to the lining
(endometrium) and burrows in
(implantation).
 Trophoblast cells send out
projections (chorionic villi) to
increase connection to maternal
blood.
Figure 38.8 A Human Blastocyst at Implantation (A) The mammalian blastocyst consists of an inner cell mass adjacent to a fluid-
filled blastocoel and surrounded by trophoblast cells. (B) Molecules and enzymes secreted by trophoblast cells allow the blastocyst
to adhere to and burrow into the endometrium. Once the blastocyst is implanted in the uterine wall, the trophoblast cells send out
chorionic villi—projections that increase the embryo’s area of contact with the maternal bloodstream.
Gastrulation Produces a
Second, then a Third
Germ Layer
38.3
Gastrulation Produces a Second, then
a Third Germ Layer

 Gastrulation involves movement of cells in the embryo.

 The three germ layers of triploblastic animals are formed; the embryo becomes
a gastrula.
Gastrulation Produces a Second, then
a Third Germ Layer

 In sea urchins (little yolk, radial cleavage):

 Vegetal pole invaginates and forms the archenteron, or gut; opening is the
blastopore

 Outer layer of cells is now ectoderm, and archenteron wall is endoderm and
future mesoderm
 Gastrulation Produces a Second, then
a Third Germ Layer

 Amphibian eggs (moderate amount of yolk):

 Cells in the gray crescent invaginate to form a blastopore.

 Cells from the animal pole begin to roll like a sheet over the dorsal lip of the
blastopore and push into the blastocoel.

 Animal hemisphere cells grow over the other cells.

Figure 38.10 Gastrulation in a Frog
Embryo
Gastrulation in Birds and Reptiles

 Birds and reptiles (eggs with lots of yolk):

 Blastopore forms as a groove in the primitive streak, a collection of cells with
Hensen’s node at the anterior end.

 Cells that will become mesoderm and endoderm migrate inward along the
streak.
Gastrulation

 Placental mammals retain the bird gastrulation pattern.

 The inner cell mass splits into an upper layer (epiblast) and a lower layer
(hypoblast).
 Embryo forms from the epiblast; hypoblast contributes to the
extraembryonic membranes and placenta.
Gastrulation

 All three types of gastrulation are found in both protostomes and

deuterostomes.
 In protostomes, the blastopore becomes the mouth and the anus forms at
some distance away.
 In deuterostomes, the blastopore becomes the anus, and the mouth
breaks through secondarily.
Gastrulation

 Gastrulation results in three

germ layers and sets the
stage for formation of the
coelom, a space completely
enclosed in mesoderm.
 In deuterostomes, the
mesoderm often forms from
the archenteron wall, and the
coelom develops within in it.
Concept 38.3 Gastrulation Produces a
Second, then a Third Germ Layer

 In protostomes, the pattern fits with mosaic development:

 Divisions of one blastomere form the mesoderm.

 The coelom forms as a split in the prospective mesoderm cells.

Gastrulation Sets the
Stage for Organogenesis
and Neurulation in
Chordates
38.4
Organogenesis and Neurulation in
Chordates

 Organogenesis: developmental phase when all organs and organ systems

form simultaneously
 Neurulation: formation of the nerve cord and brain
 The three germ layers give rise to defined tissues and organs.
 Organogenesis and Neurulation in
Chordates
 Ectoderm forms epidermis of the
skin and related structures, brain,
and nervous system.

 Endoderm forms lining of the

digestive tract, lungs, liver,
pancreas, and gall bladder.

 Mesoderm gives rise to the

notochord, heart, blood and blood
vessels, urogenital system, muscles,
bones, and the dermis (inner layer
of the skin).
Organogenesis and Neurulation in
Chordates

 When a cell displays its final specialized characteristics, it is said to be

differentiated.
 Differentiation takes place gradually throughout development and often
into adult life.
 The Notochord Induces Formation of
the Neural Tube
 In chordates, some of the cells that move through the blastopore during
gastrulation form the notochord.
 Notochord provides structural support and induces overlying ectoderm to
form the dorsal nerve cord.
 Induction: one tissue directs another tissue along a developmental
pathway by chemical signaling
The Notochord Induces Formation of
the Neural Tube

 The ectoderm over the notochord

flattens and rolls up to form the neural
tube.
 At the anterior end, three swellings in
the neural tube will become the
hindbrain, midbrain, and forebrain.
 The rest of the neural tube will become
the spinal cord.
Figure 38.13 Gastrulation, Mesoderm Formation,
and Neurulation in the Lancelet Amphioxus, a Non-
vertebrate Chordate (Part 1)
Figure 38.13 Gastrulation, Mesoderm Formation,
and Neurulation in the Lancelet Amphioxus, a Non-
vertebrate Chordate (Part 2)
The Notochord Induces Formation of
the Neural Tube

 The transcription factor β-catenin plays a key role in determining which

cells become the organizer.
 It triggers a complex series of interactions between transcription factors
and growth factors that control gene expression.
The Notochord Induces Formation of
the Neural Tube

 Development of the amphibian nervous system

 Polypeptides (Noggin and Chordin), released from the notochord, induce

ectoderm cells to become neuronal.
 They inhibit activity of the TGF-β bone morphogenetic protein BMP4.
 BMP4 induces the ectoderm to form epidermis. In the absence of BMP4,
ectoderm forms neural tissue.
The Notochord Induces Formation of
the Neural Tube

 Another inducer released by the notochord is the transcription factor Sonic

hedgehog (Shh).
 It directs the development of the ventral region of the neural tube into the
ventral structures and circuits of the spinal cord.
The Notochord Induces Formation of
the Neural Tube

 As the neural tube is closing, certain cells dissociate from it and come to lie
between the neural tube and overlying epidermis.
 These neural crest cells are pluripotent—they can differentiate into many cell
types.
 They form sensory neurons and major parts of the autonomic nervous
system, most of the skull bones, pigment cells, and many other structures.
Figure 38.16 Neurulation and
Differentiation of Mesoderm in
Vertebrates (Part 1)
Figure 38.16 Neurulation and
Differentiation of Mesoderm in
Vertebrates (Part 2)
Figure 38.16 Neurulation and
Differentiation of Mesoderm in
Vertebrates (Part 3)
The Notochord Induces Formation of
the Neural Tube

 On either side of the notochord,

three regions of mesoderm can
be recognized—somites,
intermediate mesoderm, and
lateral plate mesoderm.
 Body segments are first seen as
the somites. They form bones,
cartilage, skeletal muscle, and
dermis of the skin.
The Notochord Induces Formation of
the Neural Tube

 Intermediate mesoderm forms the urinary and reproductive system.

 Lateral plate mesoderm surrounds the coelom and lines the vertebrate
peritoneal cavity.
 Inner layer of lateral plate mesoderm forms part of the peritoneum,
muscles of the digestive tract, most of the circulatory system, including the
heart.
 Outer layer forms part of the peritoneum and some muscles of the body
wall.
Done!
2 NEXT SECTIONS ARE OUT OF SCOPE!
Extraembryonic
Membranes Protect and
Nourish the Embryo
38.5
Concept 38.5 Extraembryonic
Membranes Protect and Nourish the
Embryo
 Mammals and reptiles including birds have amniotic eggs, which provide
the embryo with a contained aqueous environment.
 Four extraembryonic membranes develop from the germ layers and
surround the embryo.
 They function in protection, nutrition, gas exchange, and waste removal.
Concept 38.5 Extraembryonic
Membranes Protect and Nourish the
Embryo
 In the chick embryo, four membranes form:
• Yolk sac—encloses yolk; can form blood vessels

 Yolk is digested by the endoderm of the yolk sac and transported to the embryo by the
blood vessels in the yolk sac wall.

• Allantois—sac for waste storage; also has blood vessels

Concept 38.5 Extraembryonic
Membranes Protect and Nourish the
Embryo
 Amnion—surrounds the embryo and secretes fluid into the enclosed cavity,
providing a protective, shock-absorbing environment

 Chorion—just beneath the egg shell; limits water loss; fuses with the allantois to
form a membrane for exchange of O2 and CO2
Figure 38.18 The Extraembryonic
Membranes of Amniotes (Part 1)
Figure 38.18 The Extraembryonic
Membranes of Amniotes (Part 2)
Figure 38.18 The Extraembryonic
Membranes of Amniotes (Part 3)
Concept 38.5 Extraembryonic
Membranes Protect and Nourish the
Embryo
 In placental mammals, hypoblast cells proliferate to form what in birds
would be the yolk sac.
 The allantois and chorion combine, forming the chorioallantoic placenta,
which combines with the endometrium to form the placenta.
 The placenta is unique because is contains tissues from two organisms—
the mother and the fetus.
Figure 38.19 The Mammalian
Placenta
Concept 38.5 Extraembryonic
Membranes Protect and Nourish the
Embryo
 Human gestation is divided into trimesters of about 12 weeks each.
 The first trimester is a time of rapid cell division and tissue differentiation
and the embryo is very sensitive to damage from radiation, drugs,
chemicals, and pathogens.
 By the end of the first trimester, most organs have started to form and the
embryo becomes a fetus.
Concept 38.5 Extraembryonic
Membranes Protect and Nourish the
Embryo
 Many fish have embryonic yolk sacs.
 As the embryo forms, all three germ layers grow around the yolk.
 The yolk sac becomes vascularized and materials are carried in the blood
vessels to the embryo.
Figure 38.20 Fish Yolk Sac
Concept 38.6 Development Continues
throughout Life

 Development does not stop at birth or hatching.

 Some animals stop growing at adulthood, some keep growing throughout
their lives.
 Direct development: animal immediately looks like the parent and grows
to reach full size
 Indirect development: life stages have very different forms
Development Continues
throughout Life
38.6
Concept 38.6 Development Continues
throughout Life

 Allometry: change in proportions of body parts relative to one another

due to unequal growth
 Isometric growth is a 1:1 size increase.
 Allometry is common in direct developers, such as humans.
 Human head to trunk ratio changes; the baby’s head is relatively larger.
Figure 38.21 Allometric Growth of a
Human
Concept 38.6 Development Continues
throughout Life

 Indirect development can involve drastic changes in morphology.

 Larval forms are often quite unlike adult forms; for example, caterpillars
and butterflies.
 Larval and adult stages may have different functions:
 Larvae can be the main feeding stage.
 Some adult insects do not feed at all and are specialized for reproduction.
Figure 38.22 Larval Forms Are Often
Quite Unlike Their Parents
Concept 38.6 Development Continues
throughout Life

 Larvae can be important in dispersal, especially if adults are sessile or

sedentary.
 Example: Corals—new sites are colonized by swimming larvae.
Concept 38.6 Development Continues
throughout Life

 Metamorphosis: major portions of the body may be remodeled or even

discarded

 Example: development of amphibian eggs into larvae (tadpoles) and then

adults.

 The tadpole undergoes changes in every system—tail and gills are resorbed,
limbs grow, gut changes animal switches from herbivory to carnivory.