Reproduction and

Development
Chapters 7-8
Asexual versus Sexual
Reproduction
 Asexual Reproduction
 Reproduction without
gametes
 No special
reproductive organs
or cells
 Only 1 parent
(usually)
 Offspring are
genetically identical
 Simple and fast
 Sexual Reproduction
 Specialized reproductive organs and cells
 Parents produce gametes (germ cells) through meiosis
 (Define germ cell and somatic cell)
 Gametes unite (fertilization) to form zygote
 2 parents (usually)
 Sexual Reproduction
 Offspring are genetically unique
Genetic material received from two
unique individuals (parents).
Sexual Reproduction (cont.)
 Result is greater genetic variation
 Evolutionary potential may be greater due to
the large number of ways genetic material
may be combined
 Sexual Reproduction (cont.)
 But process is slower
and more complex, (and
dangerous) than
asexual reproduction

Male-Male Competition for Mates Decapitation of ♂ by ♀ Preying Mantis

 Sexual Reproduction
 Why Sex if Asexual Reproduction is
easier and faster ??

 Many theories, but it appears that

in
the long run sexual organisms win
out over asexual organisms due to
genetic variation.
 ASEXUAL REPRODUCTION
 Genetically identical offspring are called
clones
 4 types
 Fission

 Budding

 Gemmulation

 Fragmentation
 Asexual Reproduction
 Fission (=dividing into
parts)
 Parent divides into 2 or
more equal parts through
mitosis
 “Binary fission” is
division into 2 equal
parts; common in
bacteria and protozoa
 Asexual Reproduction
 “Multiple fission” involves division
of the cell nucleus multiple times,
and then division of the parent into
multiple parts; Example: spore
formation (sporogony) in some
parasitic protozoa
 Asexual Reproduction
 Budding
 Offspring are
produced as an
outgrowth (bud)
which subsequently
detaches itself
 Common in
cnidarians (jellyfish)
 Asexual Reproduction
 Gemmulation
 Parent encapsulates a clump of cells in a
protective “gemmule”  (”gem-yul”)
 These cells survive freezing and/or desiccation that
kills the parent
 In the spring, the protected cells emerge from the
gemmule and produce new organisms
 Common form of reproduction in sponges
A gemmule from a freshwater sponge. 
The calcareous spicules projecting from the covering
of the gemmule and provide additional protection.
 Asexual Reproduction
 Fragmentation
 Parent breaks into 2
or more parts, and
each part grows into
a new individual
 Common in many
invertebrates like
Planarians
 SEXUAL REPRODUCTION
 Bisexual Reproduction
 2 parents
 “Dioecious” - Each has either male or female
reproductive organs, but not both.
 Unisexual Reproduction
 1 parent; 2 types
 “Monoecious” - Organism has both male and
female reproductive organs within the same
body; also called “hermaphroditism”
 “Parthenogenesis” - Production of offspring
via unfertilized eggs
Both Asexual and Sexual cycles
occur in some organisms.
 SEXUAL REPRODUCTION

Bisexual Reproduction Unisexual Reproduction

Monoecious
“=One house”

hermaphroditism

Dioecious
“=Two house”

Earthworms
 Hermaphroditism
 Common in sessile, burrowing, or
endoparasitic invertebrates
 Examples: flatworms, segmented worms,
barnacles, snails, some fishes
 Most do not fertilize themselves
 Unique form, “Sequential hermaphrodites”
change sex as they age; occurs in reef fishes
 Juvenile-not sexually active

Initial phase-
Usually female

Terminal phase-
Always a male
 Parthenogenesis
 Occurs in invertebrates as well as fish,
amphibians, reptiles, and even some turkey
strains
 6 of the 11 species of whiptails (Cnemidophorus
sp.) that occur in Texas are parthenogenetic
 Sapotes
Weevils
Described from the
Chihuahuan desert

Both
Parthenogenic
 Sexual Reproduction
 Gonads - organs that produce germ cells
(gametes)
 Gonads are termed “primary sex organs”
 “Accessory sex organs” are those that are used
in reproduction but do not produce gametes
(e.g., penis, vagina, uterus)
 Sexual Reproduction
 Males
 Gonad is the testis
 Testis produces spermatozoon (sperm)

 Sperm are small, motile, produced in large

numbers
 Females
 Gonad is the ovary
 Ovary produces ova (eggs)

 Ova are large (due to stored yolk), nonmotile,

produced in small numbers
 Sex Determination in Humans
 Do you remember what makes a male?
 Chromosomes?
 Primary sex organs?

 Accessory sex organs?

 Behavior?
 Sex Determination in Humans
 Gonads are initially the same
 In males,
 the SRY gene (sex-determining region Y) causes
development of gonad into a testis
 the testis produces testosterone

 testosterone and dihydrotestosterone (DHT) masculinize

the fetus (development of penis, scrotum, etc.)
 testosterone is converted into estrogen in the brain;
estrogen causes development of male-typical behaviors
 Sex Determination
(Experiment with Rabbits)
Fetus Fetus
(Xy) (Xy)

testosterone and (DHT) Production of

produced testosterone and (DHT)
blocked

Male Individual Female Individual

 Gamete Formation
 Spermatogenesis - produces sperm
 Oogenesis - produces ova (eggs)
Origin and Maturation of Germ Cells

A. Germ Cells
1. Somatic cells are non-reproductive body cells;
they differentiate, function and die before or with
the animal
2. Germ cells form gametes; the germ cell line provides
a continuous line between generations.
3. Somatic cells support, protect and nourish the
germ cell line.
4. The germ cell lineage may be traceable;
in some invertebrates, the germ cells develop
from somatic cells.
Migration of mammalian primordial
germ cells
B. Migration of Germ Cells

1. Vertebrate gonads arise from a pair of genital ridges

that grows into the coelom from the dorsal
coelomic lining on each side of the hindgut near
the anterior end of the kidney.
2. Primordial germ cells themselves arise from yolk-sac
endoderm, not the developing gonad.
3. Germ plasm from the vegetal pole of the uncleaved
egg mass moves to gut endoderm and migrates by
ameboid movement to genital ridges.
4. Germ cells divide first by mitosis, increasing from
a few dozen to several thousand.
 Male Reproductive System

 Testes - male reproductive organs

 Scrotum - a pouch of skin that encloses the
testes
 Temperature inside the scotum is about 34 C
 Lower temperature required for sperm production
and survival
 Muscles elevate or lower the testes depending on
air temperature
 Male Reproductive System

 Within each
testis
 Seminiferous
tubules -
produce sperm
 Male Reproductive System
 Interstitial
(Leydig) cells -
secrete
testosterone

These occur in the

connective tissue
between the
seminiferous tubules
 Spermatogenesis
 Each spermatocyte undergoes meiosis to form
4 haploid spermatids, which develop into
mature sperm (spermatozoa)
 Sperm consists of
 head with nucleus (contains chromosomes) and
acrosome (contains lysins for penetrating the egg)
 midpiece with mitochondria

 tail consisting of a flagellum

 Male Reproductive System
 From the testis,
sperm is carried
through the
 Epididymis (Develop
motility)
 vas deferens (ductus
deferens)
 Male Reproductive System
 Secretions that
provide nutrients and
lubrication are added
at ejaculation by
 seminal vesicles
 prostate gland

 bulbourethral glands
 Male Reproductive System
 Sperm plus these glandular secretions is called
“semen”
 Average human ejaculation produces 3-4 ml of
semen and 300-400 million sperm
 Semen is transported through the urethra in the
penis during ejaculation
 Urine is carried through the urethra during
urination
 Male Reproductive System
 The prepuce or foreskin is skin that folds over
the tip of the penis
 Circumcision is the removal of the prepuce or
foreskin
 Female Reproductive System
 Ovaries - female reproductive organs. They produce
estrogen and progesterone
 Oviducts (uterine tubes, fallopian tubes) - carry eggs
from ovary to the uterus. Fertilization occurs in upper
third.
 Uterus - houses the developing embryo
 Vagina - receives semen from the penis during
intercourse
 Vulva - external genitalia (labia, clitoris)
Produces estrogen and progesterone
 Oogenesis
 Each oocyte undergoes meiosis, but forms
only a single haploid ovum
 Primary oocyte undergoes first meiotic division
in ovary, produces secondary oocyte and polar
body
 Secondary oocyte is released from follicle at
ovulation
 If secondary oocyte is penetrated by sperm,
second meiotic division occurs producing
second polar body and ovum
 Sperm nucleus merges with the oocyte nucleus
to form a zygote
 Hormonal Control of
Reproduction
 Ovaries produce
 Estrogen (several forms)
 progesterone
 Testes produce
 testosterone
 DHT
(dihydrotestosterone)
Endocrine Events that Orchestrate Reproduction

Hormonal Control of Timing of Reproductive Cycles

1. Vertebrate reproduction is seasonal or cyclic to align

with food supply and survival of young.
2. Sexual cycles are controlled by hormones.
3. The hypothalamus region of the forebrain regulates
the release of anterior pituitary gland hormones, which
stimulate tissues of the gonads.
4. Estrous Cycles
a. Females are receptive to males only during brief periods
of estrus or “heat.”
b. The estrous cycle ends with uterine lining reverting to
original state; there is no menstruation.
5. Menstrual Cycles
a. This cycle occurs in monkeys, apes and humans.
b. Females are receptive to males throughout the cycle.
c. At the end of the menstrual cycle the endometrium
(uterine lining) is discharged.
 Hormonal Control of
Reproduction
 Pituitary produces:
 FSH (follicle stimulating hormone)
 LH (luteinizing hormone)

 These hormones act on the gonads and are

classified as “gonadotropins”
 Hypothalamus produces:
 GnRH (gonadotropin releasing hormone)
endometrium
The Menstrual Cycle

1. The ovary has two phases: follicular and luteal.

2. The uterus has three phases: menstrual, proliferative
and secretory.
3. Menstruation, shedding of the uterine lining, signals
the menstrual phase.
4. The follicular phase of ovary is also occurring.
a. By day three of the menstrual cycle, blood levels of
FSH and LH rise slowly, prompting some ovarian
follicles to grow and secrete estrogen.
b. As estrogen increases, the uterine endometrium
heals and begins to thicken.
c. Uterine glands within the endometrium enlarge in
the proliferative phase of uterus.
 d. By day 10, most ovarian follicles degenerate
(become atretic) leaving one, two or three to
continue ripening.
e. Final mature follicle is the Graafian follicle; it
secretes more estrogen and also inhibin.
f. At day 13 or 14, high levels of estrogen from the
Graafian follicle stimulate a surge in GnRH from
hypothalamus.
g. This stimulates a surge of LH and some FSH from
anterior pituitary.
h. The LH surge causes the largest follicle to rupture
and release an oocyte (ovulation).
5. The luteal phase of the ovary is named for the corpus
luteum, which is the remainder of the ruptured follicle.
a. The corpus luteum responds to LH and secretes
progesterone.
 b. Progesterone stimulates the uterus to undergo
maturation and prepare for gestation.
c. If an embryo implants, the uterus enters the secretory
phase.
d. If fertilization does not occur, the corpus luteum
degenerates and hormones are no longer secreted.
e. The uterus depends on progesterone and estrogen to
maintain uterine lining; declining levels start
endometrium degeneration and lead to menstrual
discharge.
[Link] feedback among the hypothalamus, anterior
pituitary and ovary control the cycle.
[Link] is due to high levels of estrogen causing a
surge in GnRH, LH and FSH; such positive feedback is
rare since it moves events away from stable set points.
 Who controls who?
Hormonal Regulation
Hormone Source
Gonadotropin releasing hypothalamus (Brain)
hormone (GnRH)
Controls
The Pituitary Gonadotropins: pituitary gland (Brain)
Follicle Stimulating Hormone
(FST)
Luteinizing Horman (LH)
Controls
Estrogens and Progesterone: Ovaries
Controls
> Development of Secondary Sex
Characteristics in female.
> Preparation of uterus for embryo.
 Hormonal Control of Female
Reproduction
 GnRH promotes secretion of LH and FSH
 FSH promotes development of the oocyte in the follicle
of an ovary
 The follicle produces estrogen and inhibit
 estrogen rebuilds uterus
 inhibin inhibits FSH production

 Elevated estrogen levels eventually trigger a surge in LH

 The LH surge causes release of the oocyte (ovulation)
 Hormonal Control of Female
Reproduction
 The follicle that the oocyte is released from is
called the corpus luteum.
 The corpus luteum secretes estrogen and
progesterone for 10-14 days. These get the
endometrium of the uterus ready for
implantation.
 If the oocyte is not fertilized, the corpus
luteum breaks down.
 Hormonal Control of Female
Reproduction
 Without estrogen and progesterone, the
endometrium breaks down, and menstruation
occurs.
 Low progesterone and estrogen allow
secretion of FSH, and beginning of new cycle.
Hormones of Human Pregnancy and Birth

[Link] normally occurs in the outer third of the

uterine tube (ampulla).
[Link] the zygote travels to uterus, it divides by mitosis to form
a blastocyst.
[Link] about six days, on contact with the uterine lining, it buries
in the endometrium (implantation).
[Link] spherically-shaped trophoblast contains three layers:
amnion, chorion and embryo proper.
[Link] chorion is a source of human chorionic gonadotropin
(hCG); it stimulates the corpus luteum to produce
estrogen and progesterone.
6. The placenta is formed between the trophoblast and uterus.
a. The placenta is an endocrine gland, secreting hCG,
estriol, and progesterone.
b. The placenta also transfers nutrients and wastes
between mother and fetus.
c. After a month, the corpus luteum degenerates and
the placenta itself holds the lining by progesterone
and estrogen. (Figure 7.17)
7. Preparation of mammary glands to secrete milk
requires two additional hormones.
a. Prolactin (PRL) is produced by the anterior pituitary,
but is inhibited in non-pregnant women.
b. During pregnancy, elevated progesterone and
estrogen depress inhibition and PRL appears in
the blood.
c. PRL is also produced by the placenta during pregnancy.
d. Human placental lactogen (hPL) aids PRL in preparing
the mammary glands for secretion.
e. Together with maternal growth hormone, hPL stimulates
an increase in nutrients in the mother.
f. The placenta also secretes -endorphin and other
endogenous opioids that regulate appetite and mood
during pregnancy.
 g. The placenta later synthesizes peptide hormone
relaxin to allow expansion of pelvis by flexibility of
pubic symphysis.
h. Relaxin also dilates the cervix in preparation for
delivery.
[Link] or parturition begins with rhythmic contractions of
uterus called labor.
a. Placental corticotropin-releasing hormone (CRH)
appears to initiate the birth process.
b. Estrogen secreted before birth stimulates contractions.
c. Progesterone levels, which inhibit contraction, decline.
d. Prostaglandin hormones increase, making the uterus
more irritable.
e. Uterine stretching causes neural reflexes to stimulate
secretion of oxytocin from the posterior pituitary.
f. Oxytocin stimulates uterine smooth muscle
contractions.
 g. Childbirth (Figure 7.18)
1) First stage: the cervix enlarges and the
amniotic sac will rupture.
2) Second stage: the baby is forced out of the
uterus and through the vagina.
3) Third stage: the placenta or afterbirth is expelled.
h. Milk Production
1) Milk production is triggered when infant sucks
on the mother’s nipple.
2) Stimulation leads to reflex release of oxytocin
from the pituitary.
3) Oxytocin causes contraction of smooth muscles
lining ducts of mammary glands.
4) Suckling also stimulates release of prolactin,
which continues milk production
Multiple Births (Figure 7.19)

[Link] mammals are multiparous, giving birth to many

offspring at one time.
2. Some give birth only to one at a time; they are uniparous.
[Link] occur; the armadillo gives birth to four young,
all male or all female, derived from one zygote.
[Link], or identical, twins are derived from one
zygote; they have identical genomes.
[Link], dizygotic or nonidentical, twins are from two
zygotes and may not resemble each other any more
than other siblings.
6. Identical Twins
a. They may separate early and have separate placentas.
b. Two-thirds share a placenta and splitting occurred after
formation of the inner cell mass, but most have
individual amniotic sacs.
c. A few share one amniotic sac and a single placenta;
separation of the zygote occurred after day 9 of
pregnancy when the amnion has formed; these twins risk
becoming conjoined (Siamese twinning).
Figure 07.19b
Fertilization
A. Initial Event
1. Fertilization is the union of male and female gametes.
a. Fertilization provides for recombination of paternal and
maternal genes, restoring the diploid number.
b. Fertilization activates the egg to begin development.

B. Oocyte Maturation
1. Contrast with sperm.
a. The sperm eliminates nearly all cytoplasm and
condenses its nucleus.
b. The egg grows in size by accumulating yolk; it also
contains much mRNA, ribosomes, tRNA and elements
for protein synthesis.
2. Morphogenetic determinants direct the activation and
repression of specific genes in the egg.
 3. The egg nucleus grows in size, bloated with RNA and
is called the germinal vesicle.
4. Preparations in the egg occur during the prolonged
prophase.
5. After meiosis resumes, the egg is ready to fuse its
nucleus with the sperm nucleus.

C. Fertilization and Activation

1. A century of research has been conducted on marine
invertebrates, especially sea urchins.
2. Contact and Recognition Between Egg and Sperm
a. Marine organisms release enormous numbers of sperm
in the ocean to fertilize eggs.
b. Many marine eggs release a chemotactic molecule to
attract sperm of the same species.
c. Sea urchin sperm first penetrate the jelly layer before
contacting the vitelline envelope.
 d. Egg-recognition proteins on the acrosomal process
bind to species-specific sperm receptors on the
vitelline envelope.
e. In the marine environment, many species may be
spawning at the same time.
f. Similar recognition proteins are found on sperm of
vertebrate species.

3. Prevention of Polyspermy
a. A fertilization cone forms where the sperm contacts
the vitelline membrane.
b. Important changes in the egg surface block entrance to
any additional sperm.
c. Polyspermy, the entry of more than one sperm, would
cause a triploid nucleus.
d. In the sea urchin, an electrical potential rapidly spreads
across the membrane; this is the “fast block.”
 e. This is followed by the cortical reaction.
1) Thousands of enzyme-rich cortical granules
below the egg membrane fuse with the membrane.
2) The cortical granules release contents between the
membrane and vitelline envelope.
3) This lifts the envelope and forms a moat.
4) One cortical granule enzyme causes the vitelline
envelope to harden, becoming the fertilization
membrane.

4. Fusion of Pronuclei and Egg Activation

a. After sperm and egg membranes have fused, the sperm
disconnects from its flagellum.
b. The enlarged sperm nucleus is the male pronucleus and
migrates inward to contact the female pronucleus.
c. Fusion forms a diploid nucleus.
 1) Nuclear fission takes 12 minutes in sea urchins; about
12 hours in mammals.
2) The fertilized egg is now properly called a zygote.

d. Fission removes one or more inhibitors that blocked

metabolism and kept the egg quiescent.
e. DNA and protein synthesis undergoes a burst of activity,
using a supply of mRNA in the egg cytoplasm.
f. Fertilization initiates reorganization of cytoplasm and
repositions determinants that begin development and
cleavage.
Cleavage and Early Development

A. Blastomeres
1. The embryo undergoes cleavage to convert the large
cytoplasmic mass into small maneuverable cells.
2. No cell growth occurs, only subdivision until cells reach
regular somatic cell size.
3. At the end of cleavage, polychaete worms have 1000
cells, amphioxus has 9000, and frogs have 700,000.
4. Polarity—a polar axis—establishes the direction of
cleavage and differentia

B. Patterns of Cleavage
1. The pattern of cleavage is affected by:
a. quantity and distribution of yolk present, and
b. genes controlling the symmetry of cleavage.
 2. The yolk-rich end of the embryo is the vegetal pole
while the other end is the animal pole.
3. Cleavage is an orderly sequence of cell divisions.
4. During each division, a cleavage furrow is visible in
the cell.

[Link] Amount and Distribution of Yolk Affects

Cleavage
1. Isolecithal yolk describes eggs with very little yolk and
the yolk is distributed evenly.
a. In such eggs, cleavage is holoblastic.
  b. The cleavage furrow extends completely through the
egg.
c. Isolecithal eggs are widespread and seen in:
echinoderms, tunicates, cephalochordates, molluscs
and mammals.
d. Cleavage is slowed in the yolk-rich vegetal pole.
2. Mesolecithal eggs have a moderate amount of yolk
concentrated in the vegetal pole.
a. The animal pole is opposite the vegetal pole and
contains cytoplasm and very little yolk.
b. These eggs cleave holoblastically, but cleavage is
retarded in yolk-rich vegetal pole.
c. The cleavage furrow progresses much more slowly
through the vegetal pole; thus cleavage is faster in
the animal region.
d. Amphibians have mesolecithal eggs.
3. Telolecithal eggs have much yolk concentrated at the
vegetal pole.
a. Actively dividing cytoplasm is confined to a narrow
disc-shaped mass on the yolk.
b. Cleavage is partial or meroblastic; the furrow does
not cut through the heavy yolk.
c. Birds, reptiles, most fishes and a few amphibians
have telolecithal eggs.
4. Centrolecithal eggs have a large mass of centrally located
yolk.
a. Cytoplasmic cleavage is limited to a surface layer of yolk-
free cytoplasm; yolk-rich inner cytoplasm is uncleaved.
b. They have meroblastic cleavage.
c. Insects and many other arthropods have centrolecithal
eggs.
d. Yolk is therefore an impediment to cleavage.
5. Amount of yolk affects developmental mode.
a. In most animals, a mother does not directly nourish
embryonic development but has provisioned the egg
with yolk.
b. The amount of yolk is related not only to cleavage pattern,
but also to whether a larval stage occurs during
development.
c. Animals in which the zygote is telolecithal generally have
direct development.
d. Species with isolecithal or mesolecithal zygotes generally
have indirect development.
e. Indirect development is characteristic of animals where
the larval stage is between embryo and adult.
f. Direct development can occur when there is enough yolk
to support growth as juveniles; this occurs in reptiles
and birds.
 If Fertilization Occurs…
 Sperm and egg nuclei fuse, becoming diploid, and
begin to divide.
 This early division is termed cleavage.
 Pattern of cleavage
 dependent on

 organism.
 Blastulation
 After mitosis has created several hundred to a
thousand cells, a ball called a blastula is formed.
 An Overview of Development Following Cleavage

A. Blastulation
1. Cleavage creates a cluster of cells called the blastula.
2. In most animals, these cells are arranged around a
fluid-filled cavity called the blastocoel.
3. The blastula stage typically consists of a few hundred
to several thousand cells.
4. Blastula formation, with a single germ layer, occurs in all
multicellular animals. In all animals other than
sponges,
development continues to form one or two more
germ layers.
5. The germ layers formed during development ultimately
produce all structures of the adult body.
 Gastrulation
 The blastula further grows and forms a gastrula.
 In more advance animals, the differentiation of
tissues begins.
 B. Gastrulation and Formation of Two Germ Layers

1. Gastrulation results in the formation of a second

germ layer
2. Typically, gastrulation involves an invagination of one
side of the blastula.
3. This invagination forms a new internal cavity,
the archenteron or gastrocoel.
4. The opening into the cavity is the blastopore.
5. The gastrula has an outer layer of ectoderm and an
inner
layer of endoderm. The only opening into this
embryonic
gut is at the blastopore.
6. Some animals retain a blind, or incomplete gut,
but most have a complete gut with a second opening,
the anus.
 C. Formation of a Complete Gut

1. In the formation of a complete gut, the inward

movement
of the archenteron continues until the end of the
archenteron reaches the ectodermal wall of the
gastrula.
2. An endodermal tube, the gut, is surrounded by the
blastocoel.
3. The endodermal tube has two openings, the
blastopore
and a second opening formed by the merging of the
archenteron tube with the ectoderm.
 Germ Layers

 If differentiation
occurs, there may be up
to three layers.
 Ectoderm – outer layer
 Mesoderm – middle
layer
 Endoderm – inner layer
D. Formation of Mesoderm, a Third Germ Layer

1. Animals with two germ layers are diploblastic.

2. Most animals add a third germ layer and are triploblastic.
3. The third layer, mesoderm, lies between the endoderm
and the ectoderm.
4. Mesoderm can form in two different ways: the proliferation
of cells from near the lip of the blastopore into the space
between the archenteron and the outer body wall,
or by the pushing of the central region of the archenteron
wall into the space between the archenteron
and the outer body wall.
5. Regardless of method of mesoderm formation, initial
mesoderm cells arise from endoderm.
 E. Formation of the Coelom

1. A coelom is a body cavity surrounded by mesoderm.

2. The coelomic cavity appears within the mesoderm by
either schizocoely or enterocoely.
3. The method by which the coelom forms is an inherited
character and is important in grouping organisms
based
on developmental characters.
4. Upon completion of coelom formation, the body has
three
tissue layers and two cavities.
 Differences
in
Development
Between
Groups
 1. There are two major groups of triploblastic animals,
protostomes and deuterostomes.
2. The groups are identified by a suite of four
developmental
characters.
A. Deuterostome Development
1. Cleavage Patterns
a. Radial cleavage – embryonic cells are arranged in
radial symmetry around the animal-vegetal axis.
b. After the third cleavage, an upper tier of cells sits
directly on top of the tier of cells below it.
c. Most deuterostomes have regulative development
in which the fate of a cell depends on interactions
with neighboring cells.
d. Early in the development of these embryos, each
cell
is capable of producing an entire embryo if
 2. Fate of Blastopore
a. Deuterostome embryos develop a complete gut.
b. The blastopore becomes the anus and second
opening
becomes the mouth.

3. Coelom Formation
a. In enterocoely, the mesoderm and the coelom form
at the same time.
b. In enterocoely, gastrulation begins with one side of
the
blastula bending inward to form the archenteron.
c. As the archenteron elongates, its sides push outward
and expand into a pouch-like coelomic
compartment.
d. This pouch-like compartment pinches off to form a
mesodermally bound space surrounding the gut.
4. Examples of Deuterostome Development –
Variations in Deuterostome Cleavage

a. Radial cleavage is characteristic of the Deuterostomia,

including echinoderms, hemichordates and chordates.
b. In bilateral cleavage, the egg is defined by unequal
cytoplasmic components.
c. The first cleavage furrow of bilateral cleavage passes
through the animal vegetal axis dividing the
asymmetrically divided cytoplasm between the two
blastomeres.
d. This first cleavage pattern determines the future right
and left side.
e. The half-embryo formed on one side is the mirror image
of the half embryo on the other side.
 f. Ascidians (tunicates) demonstrate this cleavage pattern.
g. In rotational cleavage, the first cleavage plane is
aligned with the animal vegetal axis.
h. However, in the second cleavage, one blastomere
divides meridionally while the other divides equatorially,
rotated 90 degrees to the first.
i. Early divisions may be asynchronous and possess odd
numbers of cells below the 2-4-8-16 series that would
occur with synchronous division.
j. After the third division, cells form a tightly packed cluster
stabilized by outer cells with tight junctions,
the trophoblast.
k. The trophoblast will form the embryonic portion of the
placenta.
l. Cells that give rise to the embryo are the inner cell mass.
m. This type of cleavage is present in mammals and is
slower than in other animal groups.
 n. Telolecithal eggs may divide by discoidal cleavage.
o. In telolecithal eggs, there is a large mass of yolk in each
egg; cleavage is confined to a small disc of cytoplasm.
p. Early cleavage furrows carve the disc into a single layer
of cells called the blastoderm.

[Link] of Deuterostome Development – Variations

in Deuterostome Gastrulation
a. In sea stars, gastrulation begins with the flattening of the
vegetal area of the blastula to form the vegetal plate.
b. The archenterons elongates toward the animal pole and
its anterior end expands to form two pouchlike
coelomic vesicles.
c. The ectoderm will give rise to the epithelium of the body
surface and to the nervous system.
d. Endoderm will give rise to the epithelial lining of the
digestive tube.
 e. The mesoderm, developed from the outpocketing of the
archenterons, will form the muscular system, the
reproductive system, the peritoneum, and the
calcareous plates of the sea star’s endoskeleton.
f. Frogs are deuterostomes with radial cleavage in which
the movements during gastrulation are influenced
by the large mass of yolk.
g. Slower cleavage in the vegetal half of the blastula results
in an asymmetry, with many small cells in the animal
half and fewer, larger ones in the vegetal half.
h. Gastrulation in amphibians begins with invagination of
cells located on the future dorsal side of the embryo
with the formation of a slitlike blastopore.
i. Gastrulation progresses as sheets of cells turn inward
and move inside the gastrula to form mesoderm and
ectoderm.
 j. The primitive streak forms the anterioposterior axis and
center of early growth in bird and reptile embryos.
k. The blastoderm consists of two layers: epiblast and
hypoblast with a blastocoel between them.
l. The epiblast sheet moves toward the primitive streak
and over the edge, migrating as cells into the
blastocoel.
m. One stream of cells moves deeper, displacing midline
hypoblast, and forms endoderm.
n. Surface cells form ectoderm.
o. All three layers lift from the yolk and pinch off, leaving a
stalk attachment to the yolk at midbody.
p. In mammalian gastrulation, epiblast cells move medially
through the primitive streak into the blastocoel; cells
migrate laterally through the blastocoel to form
mesoderm and endoderm.
 q. Endoderm derived from the hypoblast forms a yolk
sac without yolk; mammals utilize nutrients from a
placenta.
r. Reptiles, birds and mammals share a common
ancestor whose eggs were telolecithal; all inherited
this gastrulation pattern and mammals then evolved
isolecithal eggs with a telolecithal pattern

B. Protostome Development
1. Cleavage Patterns
a. Spiral cleavage occurs in most protostomes.
b. Blastomere cleave obliquely to the animal-vegetal
axis.
c. Upper layer of cells appears offset relative to lower layer.
d. Mosaic development is characteristic of most protostomes.
e. In mosaic development, cell fate is determined by the
distribution of morphogenetic determinants,
in the egg cytoplasm.
 2. Fate of Blastopore

a. Protostomes are so named because the blastopore

becomes the mouth.
3. Coelom Formation

a. In protostomes, a band of mesoderm forms around

the gut before a coelom forms.
b. If a coelom exists, it is formed by schizocoely.
c. Endodermal cells move by ingression into the space
between the archenteron walls and the ectoderm.
d. These cells divide and produce mesodermal
precursors;
the proliferating cells become the mesoderm.
e. Studies indicate that these mesodermal precursors
originate from a single large blastomere, the 4d cell,
present in a 29- to 64- cell embryo.
 f. Some protostomes, like flatworms, are acoelomate;
g. In others, mesoderm lines only one side of the
blastocoel, leaving a fluid-filled cavity around the gut
called a pseudocoelom.
h. The pseudocoelom is lined on the outer edge by
mesoderm and on the inner edge by the
endodermal gut lining.

4. Examples of Protostome Development

a. The protostomes are divided into two clades, the
lophotrochozoan protostomes and the
edcysozoan protostomes.
b. The lophotrochozoan protostomes contains
segmented worms, molluscs, a some less well-known
taxa.
c. Some members of this clade employ a whorl of
tentacles
known as a lophophore and a trochophore larva.
 d. The edcysozoan protostomes includes arthropods,
roundworms, and other taxa that molt the exoskeleton;
the name of the clade refers to ecdysis.

[Link] of Protostome Development – Variations in

Protostome Cleavage
a. Although spiral cleavage is typical of protostomes, the
cephalopod mollusks use bilateral cleavage.
b. Many ecdysozoans do not exhibit spiral cleavage; in
some, cleavage appears radial while in others it is of a
unique type such as the superficial cleavage of insects.
c. In superficial cleavage, the centrally located mass of yolk
restricts cleavage to the cytoplasmic rim of the egg.
6. Examples of Protostome Development – Variations in
Protostome Gastrulation

a. In most protostomes, mesodermal cells all derive from

the 4d cell.
b. In some nemertean worms, mesoderm derives from an
earlier blastomere.
Similarities in Development
Within Groups
Vertebrate Development

A. Common Vertebrate Heritage

1. One outcome of shared ancestry in vertebrates is the
similarity of postgastrula embryos.
a. For a short time, all vertebrate embryos share: dorsal
neural tube, notochord, pharyngeal gill pouches with
aortic arches, ventral heart and postanal tail.
b. This similarity is extraordinary considering the variety
of eggs and developmental patterns.

B. Amniotes and the Amniotic Egg

1. Amniotes are a monophyletic grouping of vertebrates.
a. Their embryos develop within the amnion, a
membranous sac that provides an aquatic environment
and protects it from shock.
 b. The amniotic egg contains four extraembryonic
membranes including the amnion.
c. The amniotic egg allowed development away from
water.
d. The yolk sac is the first membrane and pre-dates the
amniotes by millions of years; it is extraembryonic
and is discarded.
e. The allantois is a sac that grows out of the hindgut
and holds metabolic wastes during development.
f. The chorion lies beneath the eggshell and encloses
the rest of the embryo.
g. With growth, the allantois and chorion fuse to form
the chorioallantoic membrane, a provisional “lung.”
h. Evolution of the shelled amniotic egg made
fertilization a requirement before a shell was
developed.
 Amniotic Animals
 Includes reptiles, birds and mammals.

Placental Mammals

Reptiles and Birds

C. Mammalian Placenta and Early Mammalian
Development
1. Mammals inherited the amniotic egg but retained it in the
mother’s body.
a. Monotremes are examples of primitive mammals that lay
eggs.
b. In marsupials, embryos develop but do not “take root” in
the uterus; they climb out and enter an external
abdominal pouch.
2. Placental mammals represent 94% of the Class
Mammalia.
a. Evolution of a placenta required considerable
restructuring.
b. Extraembryonic membranes had to restructure to form
the placenta.
c. The maternal oviduct had to evolve to develop a uterus
to
house the embryo.
 Early Stages of Mammalian Development

a. The blastocyst travels down the oviduct toward the uterus,

propelled by ciliary action and peristalsis.
b. At about the sixth day, the human blastocyst composed
of about 100 cells contacts the endometrium.
c. On contact, the trophoblast cells proliferate rapidly and
produce enzymes to break down epithelium of the
endometrium.
d. By the twelfth day, the blastocyst is totally buried,
surrounded by maternal blood.
e. The trophoblast thickens, sending out tiny fingers of
chorionic villi.
f. The surface area of chorionic villi eventually expands to
13 square meters in the human placenta.
g. The amnion remains unchanged; the embryo floats
in this “pond.”
h. The yolk sac contains no yolk but has stem cells that
give rise to blood and lymphoid cells; they later
migrate into the developing embryo.
i. The allantois is not needed to store wastes; it
contributes to the umbilical cord.
j. The chorion forms the embryo’s side of the placenta.
k. The germinal period is the first two weeks.
l. The embryonic period includes the next eight weeks
when all major organs and body shape form; this is a
period that is sensitive to drugs that may cause
malformation in the embryo.
m. The fetal period begins when embryo becomes a
fetus at about two months; mainly a growth phase,
but the endocrine and nervous systems still develop.
Fate of Germ Layers in Mammals