CHAPTER 45

Animal Reproductive Systems

張永福
生物醫學暨環境生物學系
第一教學大樓九樓N923
07-3121101轉2701
m795003@kmu.edu.tw
1
PowerPoint Lectures for Campbell Biology, 11th ed.© 2018 Pearson Education, Inc.
 Key Concepts
45.1 Both asexual and sexual reproduction
occur in the animal kingdom
45.2 Fertilization depends on mechanisms that
bring together sperm and eggs of the
same species
45.3 Reproductive organs produce and
transport gametes
45.4 The interplay of tropic and sex hormones
regulates mammalian reproduction
45.5 In placental mammals, an embryo
develops fully within the mother’s uterus
2
Let Me Count the Ways
 Animal reproduction takes many forms
 population outlive: reproduction

Figure 45.1 What are the tiny orbs, and where are they going?
3
45.1: Both asexual and sexual reproduction
occur in the animal kingdom
Mechanisms of Asexual Reproduction
 Sexual reproduction
− Sperm
− Egg
− zygote
 Asexual reproduction
− Fission
− Budding
− Fragmentation/ Regeneration
− Parthenogenesis: mainly in invertebrates, ex
bees, ants, Komodo dragon, Hammerhead Shark,
whiptail lizards 4
Variation in Patterns of Sexual Reproduction
 Hermaphroditism
− each individual has both male and female
reproductive systems
− Any two individuals can mate, and in some species,
hermaphrodites can also self-fertilize
− Several organisms can change their sex under
certain circumstances

Figure 45.2 Reproduction

among hermaphrodites

5
 Reproductive Cycles
 Reproductive cycles:
hormones, environment
 Ovulation (a) A. uniparens females

 Some organisms can

Ovary
size
reproduce both sexually Ovulation Ovulation

or asexually, ex Daphina Progesterone

Hormone
Estradiol

 Parthenogenesis:

level
− chromosomes doubling Time

after meiosis
− fishes, amphibians, and
Behavior: Female Male- Female Male-
like like
lizards (whiptail lizards) (b) The changes in sexual behavior of A.
uniparens individuals are correlated with the
cycles of ovulation.
Figure 45.4 Sexual behavior in parthenogenetic lizards6
Sexual Reproduction: An Evolutionary Enigma
Asexual Sexual
reproduction reproduction
Female Generation Female
1
Generation
2 Male
Generation
3
Generation
4
 Sexual reproduction: offspring of varied phenotypes,
enhance reproductive success when environment
changed
 Asexual reproduction: advantageous in stable,
favorable environments
7
45.2: Fertilization depends on mechanisms that
bring together sperm and eggs of the same species
 Fertilization
− external fertilization
− internal fertilization
Ensuring the Survival of Offspring
• fewer gametes but higher survival
zygotes
• eggs with calcium- and protein-
containing shells and several
internal membranes
• Embryo develops inside the female 8
Gamete Production and Delivery
 Gonads: organs to produce gametes
 Spermatheca: sperm stored
Accessory
 cloaca gland
1 Testis

2 Vas deferens
4 Ejaculatory
duct 3 Seminal
vesicle
5 Penis and
(a) Male fruit fly claspers
1 Ovary

Figure 45.7 2 Oviduct

Spermatheca
Insect
reproductive Accessory Uterus
gland 3 Vulva
anatomy
(b) Female fruit fly 9
Concept 45.3: Reproductive organs produce
and transport gametes
Human Male Seminal
(Urinary bladder)
Reproductive vesicle
(behind
Anatomy bladder) Prostate gland
Bulbourethral gland
Urethra
Erectile tissue of penis
Scrotum Vas deferens
Epididymis
Testis

(60% volume, alkaline,

fructose, prostaglandins..) (Urinary bladder)
Seminal vesicle (Urinary duct)
(Rectum) (Pubic bone)
Erectile
Ejaculatory duct tissue
(citrate)Prostate gland Urethra Penis
Vas deferens Glans
Bulbourethral Epididymis
gland seminiferous Testis Prepuce
(neutralized tubules Scrotum
mucus) 10
 Human Female Reproductive Anatomy
Oviduct
Ovary
Uterus
(Urinary bladder)
(Rectum) (Pubic bone)
Cervix (Urethra)
Body
Vagina Clitoris
Glans
Prepuce
Major vestibular
gland
Vaginal opening
Labia minora
Labia majora
•The mammary
glands：not in
Ovaries Oviduct
the reproductive
system
Follicles
Uterus Uterine wall
Endometrium
Cervix

Vagina
Gametogenesis  Spermatogenesis
Epididymis Seminiferous Primordial germ
Key cell in embryo
tubule
Diploid (2n) Mitotic divisions
Haploid (n) Spermatogonial 2n
stem cell
Lumen Mitotic divisions
Testis Spermatogonium 2n
Mitotic divisions
Sertoli Primary
cell 2n
spermatocyte
nucleus
Meiosis I
Secondary n n
spermatocyte
Spermatids Meiosis II
Early n n n n
Neck spermatid
Tail
Plasma Midpiece Head Differentiation
membran Lumen of
Nucleus seminiferous Sperm n n n n
e
Acrosome tubule cell
Mitochondria
12
 Primordial germ
cell Mitotic
 Oogenesis Ovary

divisions Follicle
2n Oogonium
Primary oocyte
Mitotic divisions within follicle

Primary oocyte (present

2n at birth), arrested in Growing
IN EMBRYO prophase of meiosis I follicle
Completion of meiosis I
STARTING AT and onset of meiosis II
PUBERTYn
n Secondary oocyte,
First arrested at metaphase
polar of meiosis II
body Mature follicle
Ovulation, sperm entry Ruptured
follicle

Completion of meiosis II
Second Ovulated
polar n secondary
body oocyte
n Fertilized egg

Corpus luteum

Degenerating
corpus luteum
13
 Spermatogenesis differs from oogenesis in three
ways
− All four products of meiosis develop into sperm
while only one of the four becomes an egg
− Spermatogenesis occurs from adolescence

− Sperm are produced continuously without the

prolonged interruptions in oogenesis

14
Concept 45.4: The interplay of tropic and sex
hormones regulates mammalian reproduction
 Gonadotropin-releasing hormone (GnRH):
− secreted by the hypothalamus, directs the release of
FSH (follicle-stimulating hormone) and LH
(luteinizing hormone)
 FSH and LH:
− from the anterior pituitary, regulate processes in the
gonads and the production of sex hormones
 sex hormones:
− steroid hormones
− Androgen: Testosterone
− Estrogens: mainly estradiol and progesterone
− Functions: gamete production, sexual behavior, the
development of primary and secondary sex
characteristics 15
Hormonal Control of the Male Reproductive System

Hypothalamus

GnRH

Figure 45.12 Anterior

Androgen-dependent pituitary
male anatomy and
behavior in a moose
FSH LH
feedback

feedback
Negative

Negative
Sertoli cells Leydig cells

Inhibin Testosterone

Spermatogenesis
TESTIS 16
Hormonal Control of the Female Reproductive Cycles
 menstruation: endometrium shed
 two cycles of female reproduction
− ovarian cycle:
1. follicular phase: GnRH → FSH and LH →
estradiol and follicle growth → LH → ovulation
2. ovulation: secondary oocyte released
3. luteal phase: ovulation →corpus luteum →
progesterone and estradiol
− uterine (menstrual) cycle:
1. proliferative phase: endometrium thickening
2. secretory phase: nutrients secretion
3. menstrual flow phase: endometrium shedding
17
 (a) Control by hypothalamus 9 Inhibited by combination of estradiol
Hypothalamus  and progesterone
1 GnRH  Stimulated by high levels of estradiol
5
Anterior pituitary  Inhibited by low levels of estradiol
4
2 FSH LH
(b) Pituitary gonadotropins 6
in blood

LH
FSH
3 FSH and LH stimulate LH surge triggers ovulation
(c) Ovarian cyclefollicle to grow
7 8
Degenerating
Growing follicle Maturing follicle Corpus luteum corpus luteum
Follicular phase Ovulation Luteal phase
4 Estradiol secreted by growing Progesterone and estradiol
follicle in increasing amounts Secreted by corpus luteum
(d) Ovarian hormones Peak causes LH surge
in blood 5
Estradiol 10
9
Progesterone
Estradiol level very low Progesterone and estradiol promote
Thickening of endometrium
(e) Uterine (menstrual) cycle
Endometrium
Days

Menstrual flow phase Proliferative phase Secretory phase

0 5 10 14 15 20 25 18
28
 Ectopic: cells of the uterine lining migrate to
an abnormal location, ex endometriosis
Menopause
 the cessation of ovulation and menstruation

19
Menstrual Versus Estrous Cycles
 Menstrual cycles:
− humans and some other primates
− Menstruation: endometrium shed from the uterus
− Sexual receptivity is not limited to a time frame
 Estrous cycles:
− most mammals
− The endometrium is reabsorbed by the uterus
− Estrus: sexual receptivity is limited to “heat”
− The length and frequency of estrous cycles vary
from species to species

20
Human Sexual Response

 Two reactions predominate in both sexes:

− Vasocongestion: the filling of tissue with blood

− Myotonia: increased muscle tension

 The sexual response cycle has four phases:

excitement, plateau, orgasm, and resolution

21
Concept 45.5: In placental mammals, an embryo
develops fully within the mother’s uterus
Conception, Embryonic Development, and Birth
 Conception: fertilization, in the oviduct (fallopian tube)
 blastocyst
 Pregnancy: gestation
3 Cleavage
4 Cleavage
continues.
Ovary
2 Fertilization

Uterus 5 Implantation
1 Ovulation

Endometrium
22
First Trimester
 human chorionic gonadotropin (hCG):
− embryo secretes hormones
− maintains secretion of progesterone and estrogens
 trophoblast :
− the outer layer of the blastocyst
− mingles with the endometrium to form placenta
 Organogenesis: development of the body organs
 Fetus: major structures of embryo present by 8
weeks

23
 Changes occur in the mother
− Mucus plug to protect against infection
− Growth of the placenta and uterus
− Cessation of ovulation and the menstrual cycle
− Breast enlargement
− Nausea is also very common
Maternal arteries Maternal veins
Placenta
Maternal
Umbilical cord portion of
placenta
Chorionic villus,
containing Fetal
Fetal capillaries portion of
placenta
Maternal (chorion)
Uterus blood pool
Umbilical
Fetal arteriole Fetal venule arteries
Umbilical cord Umbilical vein
24
Figure 45.17

(a) 5 weeks (b) 14 weeks

Figure 45.17 Some stages of human development
during the first and second trimesters
Second and Third Trimesters
 the second trimester:
− The fetus grows and is very active
− The mother may feel fetal movements
− The uterus grows enough for the pregnancy
 the third trimester:
− the fetus grows and fills the space within the
embryonic membranes
 Labor:
− a series of strong, rhythmic uterine contractions
− regulated by prostaglandins and hormone such as
estradiol and oxytocin

26
Figure 45.18

Estradiol Oxytocin
from from fetus
ovaries and mother’s
posterior pituitary
Activates oxytocin

Positive feedback
receptors on uterus
Stimulates uterus
to contract

Stimulates
placenta to make

Prostaglandins

Stimulate more
contractions
of uterus
Figure 45.19

Placenta
Umbilical cord
Uterus
Cervix

1 Dilation of
the cervix

2 Expulsion:
delivery of the infant

Uterus
Placenta
(detaching)
Umbilical cord
3 Delivery of
the placenta
Maternal Immune Tolerance of the Embryo and
Fetus
 A woman’s acceptance of her “foreign” offspring is
not fully understood
 It may be due to suppression of the immune
response in her uterus

29
Contraception and Abortion
 Contraception:
− Preventing release of eggs and sperm: birth
control pills, Tubal ligation, Vasectomy
− Keeping sperm and egg apart: rhythm method,
Coitus interruptus (withdrawal), condom, diaphragm
− Preventing implantation of an embryo: Intrauterine
devices (IUDs)
 Abortion: the drug RU486 terminates a
pregnancy nonsurgically within the first 7 weeks

30
 MALE FEMALE
Method Event Event Method
Production of Production of
sperm primary oocytes
Vasectomy Combination
birth control
Sperm transport Oocyte pill (or injection,
down male development patch, or
vaginal ring)
duct system and ovulation
Abstinence Abstinence
Condom Female condom
Coitus
interruptus
(very high Sperm Capture of the
failure rate) deposited oocyte by the
in vagina oviduct
Tubal ligation
Spermicides;
diaphragm;
Sperm movement Transport progestin alone
through female of oocyte in (as minipill
reproductive oviduct or injection)
tract

Meeting of sperm and oocyte

in oviduct
Morning-after pill;
Union of sperm and egg intrauterine device
(IUD)
Implantation of blastocyst
in endometrium
Modern Reproductive Technologies
Detecting Disorders During Pregnancy
 Invasive: amniocentesis and chorionic villus sampling
 Noninvasive: ultrasound imaging
Infertility and In Vitro Fertilization
 In vitro fertilization (IVF): mixes eggs with sperm in
culture dishes and returns the embryo to the uterus at
the eight-cell stage
 Sperm or sperm nuclei can also be injected directly
into an oocyte

32
 Summary by Questions

1. Distinguish between asexual and sexual

reproduction
2. Explain how hermaphroditism may be
advantageous to animals that have difficulty
encountering a member of the opposite sex
3. Describe various ways in which animals may
protect developing embryos
4. Using diagrams, identify and state the
function of each component of the male and
female reproductive systems

33
5. Describe oogenesis and spermatogenesis;
describe three major differences between
them
6. Explain how the uterine and ovarian cycles
are synchronized and describe the functions
of the hormones involved
7. List the various methods of contraception,
how each works, and how effective each is
8. Describe techniques that allow us to learn
about the health and genetics of a fetus

34
 CHAPTER 46
Development in Animals
張永福
生物醫學暨環境生物學系
第一教學大樓九樓N923
07-3121101轉2701
m795003@kmu.edu.tw
PowerPoint Lectures for Campbell Biology, 11th ed.© 2018 Pearson Education, Inc.
35
 Key Concepts
46.1 Fertilization and cleavage initiate
embryonic development
46.2 Morphogenesis in animals involves
specific changes in cell shape,
position, and survival
46.3 Cytoplasmic determinants and
inductive signals contribute to cell fate
specification

36
A Body-Building Plan
 model organisms
EMBRYONIC DEVELOPMENT
Sperm
Zygote
Adult Egg
frog

1 mm

Metamorphosis Blastula

Larval 1 mm
Gastrula
stages
Tail-bud
embryo
37
Concept 46.1: Fertilization and cleavage initiate
embryonic development
Fertilization
Sperm penetrate the protective layer of the egg
−acrosomal reaction: acrosome, hydrolytic enzymes
Receptors on the egg surface bind to molecules on
the sperm surface
Changes at the egg surface prevent polyspermy, the
entry of multiple sperm nuclei into the egg
−fast block to polyspermy: depolarizes membrane
−slow block to polyspermy:
The Cortical Reaction: Ca2+   cortical granules
release outside the egg  fertilization envelope 38
Figure 46.3-5

Sperm
plasma
membrane

Sperm
nucleus Fertilization
Acrosomal envelope
process
Basal body
(centriole) Actin
filament
Sperm
head
Fused Cortical
plasma granule
membranes
Acrosome Perivitelline
Jelly coat Hydrolytic enzymes
space
Vitelline layer
Sperm-binding
receptors Egg plasma membrane EGG CYTOPLASM

Figure 46.3-5 The acrosomal and cortical reactions during sea urchin
fertilization
© 2014 Pearson Education, Inc.
Egg Activation
Ca2+  cellular respiration and protein synthesis 
Experiment Fertilization
envelope

10 sec after 25 sec 35 sec 1 min

500 μm
fertilization
Results Spreading
Point of sperm wave of Ca2+
nucleus entry

1 sec before 10 sec after 20 sec 30 sec

500 μm
fertilization fertilization
Figure 46.4 Inquiry: Does the distribution of CA2+ in an egg correlate with formation of the
fertilization envelope? 40
Fertilization in Mammals
 zona pellucida: extracellular matrix of the egg
 No fast block to polyspermy in mammals

Zona pellucida
Follicle cell

Sperm Cortical
Sperm nucleusgranules
basal body
Figure 46.5 Fertilization in mammals
41
Cleavage
 Cleavage: rapid cell division without growth
 Blastula: a ball of cells
− Blastocoel: fluid-filled cavity
− Blastomeres: many smaller cells

50 m

(a) Fertilized egg (b) Four-cell stage (c) Early blastula (d) Later blastula

Figure 46.6 Cleavage in an echinoderm embryo

42
Cleavage Pattern in Frogs
Zygote Animal

 yolk hemisphere
Cleavage furrow
 vegetal pole Vegetal 2-cell
 animal pole 0.25 mm hemisphere
Gray
stage
forming
crescent

4-cell
Animal stage
pole forming

0.25 mm
Blastocoel
8-cell
stage

Blastula
Figure 46.7-3 Cleavage in a frog embryo (cross section) 43
Cleavage Pattern in Other Animals

 Holoblastic cleavage: complete division of the

egg, occurs in species whose eggs have little or
moderate amounts of yolk, such as sea urchins
and frogs
 Meroblastic cleavage: incomplete division of the
egg, occurs in species with yolk-rich eggs, such as
reptiles and birds

44
Concept 46.2: Morphogenesis in animals involves
specific changes in cell shape, position, and survival
Morphogenesis
 Gastrulation:
− gastrula：germ layers embryo, primitive gut
(archenteron)
− embryonic germ layers
 The ectoderm forms the outer layer
 The endoderm lines the digestive tract
 The mesoderm partly fills the space between
the endoderm and ectoderm
− blastopore
 Organogenesis: the formation of organs
45
Reorganizing the animal embryo in three dimensions
Surface view:

Plane of section
Cross section: Archenteron
Blastula Blastocoel
Gastrula

Blastopore Primitive
Forming the primary cell layers of the animal body gut
Diploblast: Hydra Adult Open end of
polyp gastrovascular
cavity
Blastopore Open end Mouth
Triploblast: Planaria (protostome)
Adult worm (on underside)
Gastrovascular
cavity
Blastopore mouth
Triploblast: Sea urchin (deuterostome) Mouth
Ectoderm
Mesoderm
Larva Anus
Blastopore anus Endoderm
Gastrulation in Frogs
SURFACE VIEW CROSS SECTION
1 Animal pole
Blastocoel

dorsal lip： Dorsal lip

of blasto- Dorsal lip
in the gray pore
Blastopore
of blastopore
Early Vegetal pole
crescent, gastrula
Blastocoel
invaginate to 2
shrinking Archenteron

create the
archenteron
Blastopore Blastopore

3 Ectoderm
Blastocoel Mesoderm
remnant Endoderm

Archenteron

Future ectoderm Blastopore

Future mesoderm Late
gastrula Blastopore Yolk plug 47
Future endoderm
Gastrulation in Chicks
 embryo: epiblast + hypoblast Fertilized egg
Primitive
 Epiblast→ embryo streak
Embryo
 hypoblast→ yolk sac Yolk

 primitive streak：similar as
blastopore Primitive streak

Epiblast

Future
ectoderm

Blastocoel
Migrating Endoderm
cells Hypoblast
(mesoderm) YOLK
48
Figure 46.9

ECTODERM (outer layer of embryo)

• Epidermis of skin and its derivatives (including sweat glands, hair follicles)
• Nervous and sensory systems
• Pituitary gland, adrenal medulla
• Jaws and teeth
• Germ cells

MESODERM (middle layer of embryo)

• Skeletal and muscular systems

• Circulatory and lymphatic systems
• Excretory and reproductive systems (except germ cells)
• Dermis of skin
• Adrenal cortex

ENDODERM (inner layer of embryo)

• Epithelial lining of digestive tract and associated organs (liver, pancreas)

• Epithelial lining of respiratory, excretory, and reproductive tracts and ducts
• Thymus, thyroid, and parathyroid glands
© 2014 Pearson Education, Inc.
Gastrulation in Humans
 Human eggs have very little yolk
 A blastocyst: human blastula
− The inner cell mass: develops into embryo
− The trophoblast: contribute to implantation
and extraembryonic membranes (Chorion)
formation
 Gastrulation involves the inward movement from
the epiblast, through a primitive streak, similar to
the chick embryo

50
 1 Blastocyst reaches uterus. Endometrial epithelium
(uterine lining)
Uterus Inner cell mass
Trophoblast
Blastocoel

2 Blastocyst implants Expanding region of

(7 days after fertilization). trophoblast
Maternal
Epiblast
blood
vessel Hypoblast
Trophoblast

Expanding region of trophoblast

3 Extraembryonic membranes
start to form (10–11 days), and Amniotic cavity
gastrulation begins (13 days). Epiblast
Hypoblast
Yolk sac (from hypoblast)
Extraembryonic mesoderm cells
(from epiblast)
Chorion (from trophoblast)
4 Gastrulation has produced Amnion
a three-layered embryo Chorion
with four extraembryonic Ectoderm
Mesoderm
membranes: the amnion, Endoderm
chorion, yolk sac,
and allantois. Yolk sac
Extraembryonic mesoderm
Allantois
© 2014 Pearson Education, Inc.
Developmental Adaptations of Amniotes
 Amniotes: embryos surrounded by fluid sac
(amnion) of mammals, reptiles and birds
 Land vertebrates form four extraembryonic
membranes:
− The chorion functions in gas exchange
− The amnion encloses the amniotic fluid
− The yolk sac encloses the yolk
− The allantois disposes of waste products
Chorion
Allantois
Amnion
Yolk sac
52
Organogenesis
Neurulation
 Notochord:
− from the dorsal mesoderm, signaling molecules
secreted (induction) to form the neural plate
(ectoderm)
− disappears before birth, but contributes to parts
of the discs between vertebrae
 Neural tube: neural plate curves inward, become
the central nervous system (brain and spinal cord)

53
 Neural folds Eye Somites Tail bud
Neural Neural plate
fold

SEM
1 mm
1 mm Neural tube Neural
Neural Neural Notochord crest
fold plate
Neural Coelom cells
crest Somite
cells
Notochord
Ectoderm Archenteron
Neural Outer layer
Mesoderm crest of ectoderm (digestive
Endoderm cells cavity)
Archenteron

Neural
tube
(a) Neural plate formation (b) Neural tube formation(c) Somites
Cell Migration in Organogenesis
Eye Somites Tail bud
 Neural crest cells:
ectoderm, migrate in
the body, eventually
forming various parts
of the embryo (nerves,
parts of teeth, skull SEM
1 mm
bones, and so on) Neural tube Neural
Notochord crest
Coelom
 Somites: mesoderm, cells
Somite
form vertebrate, ribs,
muscles
Archenteron
(digestive
cavity) 55
Organogenesis in Chick and Insects
 The mechanism of organogenesis in the chick and
invertebrates is quite similar to that in the frog

Neural tube
Notochord Eye
Archenteron Forebrain
Somite
Coelom Heart
Lateral
Endoderm
fold
Mesoderm Blood
Ectoderm vessels
Somites
Yolk stalk
These layers Yolk sac
form extraembryonic Neural tube
membranes. YOLK
(a) Early organogenesis (b) Late organogenesis

Figure 46.15 Organogenesis in a chick embryo

56
The Cytoskeleton in Morphogenesis
Morphogenesis:
 cell shape changes or cell migration,
 in animals but not plants
 cell shape changes: Ectoderm
microtubules and Neural
plate
microfilaments of
Microtubules
the cytoskeleton

Actin
filaments

Neural tube
57
 convergent extension: cells rearrange to
become narrower (converge) and longer (extend)

Convergence Extension
Cells elongate and crawl The sheet of cells becomes
between each other. longer and narrower.
Figure 46.17 Convergent extension of a sheet of cells

Programmed Cell Death

 also called apoptosis
 extra neurons are removed by apoptosis
58
Concept 46.3: Cytoplasmic determinants and
inductive signals contribute to cell fate specification
 Determination: cells becomes to a particular fate
 Differentiation: specialization

Fate Mapping
Epidermis Central Epidermis
nervous 64-cell
system embryos
Notochord
Blastomeres
Mesoderm injected with dye
Larvae
Endoderm
Blastula Neural tube stage
(a) Fate map of a frog embryo (b) Cell lineage analysis in a tunicate

Figure 46.18 Fate mapping for two chordates

59
 Cell lineage in Caenorhabditis elegans

Time after fertilization (hours)

0 Zygote
First cell division

Nervous Musculature, Outer skin, Germ line

system, gonads nervous system (future
outer skin, gametes)
muscula-
ture Musculature

10
Hatching

Intestine

Intestine

Mouth Anus
Eggs Vulva

ANTERIOR POSTERIOR
1.2 mm
P granules of C. elegans :
 complexes of RNA and protein
 In the posterior end, specification of germ cell fate
 act as cytoplasmic determinants
100 μm

Cells with P granules

20 μm

1 Newly fertilized egg 3 Two-cell embryo

2 Zygote prior to first division 4 Four-cell embryo 61

Axis Formation
Dorsal
Right

Anterior Posterior

Left
Ventral
(a) The three axes of the fully developed embryo

Animal pole Point of Pigmented

Animal
sperm cortex
hemisphere
nucleus
entry Future First
dorsal cleavage
side
Vegetal
hemisphere Gray
Vegetal pole crescent
(b) Establishing the axes 62
Restricting Developmental Potential
 Totipotent: can develop into all cell types, ex zygote
Experiment
Control egg Experimental egg
(dorsal view) (side view)
1a Control group 1b Experimental
group Gray
Gray crescent
crescent

Thread

Results

Normal Belly piece Normal 63

Cell Fate Determination and Pattern Formation
by Inductive Signals
The Spemann’s organizer：
dorsal lip
Experiment Results Dorsal lip from albino gastrula
Dorsal lip of Primary embryo
blastopore

Secondary
Pigmented (induced) embryo
gastrula
(donor embryo) Primary structures:
Nonpigmented Neural tube
gastrula Notochord
(recipient embryo)
Secondary structures:
Notochord (pigmented cells)
Neural tube
(mostly nonpigmented cells)
64
Formation of the Vertebrate Limb
 pattern formation: spatial organization development
 positional information: molecular cues control
pattern formation
 apical ectodermal ridge (AER): secret fibroblast
growth factor (FGF) to promote limb bud outgrowth
 zone of polarizing activity (ZPA): secreting signal
protein Sonic hedgehog at posterior side of the bud
Anterior
Limb bud

2
AER
Digits
ZPA
Limb buds Posterior
50 μm
3
Anterior4
Apical Ventral
ectodermal Proximal Distal
ridge (AER)
Dorsal
Posterior
(a) Organizer regions (b) Wing of chick embryo 65
Figure 46.26

 The production of a forelimb or a hindlimb depends

on patterns of Hox gene expression
 cell fates in animals: BMP-4, FGF, hedgehog, Hox
Experiment
Anterior
New
ZPA
Donor Host
limb limb
ZPA bud bud
Posterior
Results

4 3

22

4 3
© 2014 Pearson Education, Inc.
Cilia and Cell Fate
 Ciliary function is essential for proper specification
of cell fate in the human embryo
 Motile cilia play roles in left-right specification
 Monocilia (nonmotile cilia) play roles in normal
kidney development

Lungs
Heart
Liver
Spleen
Stomach
Large intestine

Normal location Location in

of internal organs situs inversus 67
 Summary by Questions

1. Describe the acrosomal reaction

2. Describe the cortical reaction
3. Distinguish among meroblastic cleavage and
holoblastic cleavage
4. Compare the formation of a blastula and
gastrulation in a sea urchin, a frog, and a
chick
5. List and explain the functions of the
extraembryonic membranes

68
6. Describe the process of convergent extension
7. Describe the role of the extracellular matrix in
embryonic development
8. Describe two general principles that integrate
our knowledge of the genetic and cellular
mechanisms underlying differentiation
9. Explain the significance of Spemann’s organizer
in amphibian development
10.Explain pattern formation in a developing chick
limb, including the roles of the apical
ectodermal ridge and the zone of polarizing
activity
69