Animal Development

Development
8. Animal Development
8.1 Fertilization
Q: What is fertilization?
Combination of haploid sets of chromosomes
from two individuals into a single diploid cell,
called the zygote.
Union of male and female gametes
 The gametes (egg and sperm) are both highly
specialized cell types
 Contact of the sperm with the egg’s surface
initiates metabolic reactions within the egg that
trigger the onset of embryonic development.
Spermatozoa
 Consists of head, midpiece (body), & flagellum
(tail)
 Head encloses a nucleus, & capped by vesicle
called acrosome
Q: What is the importance of acrosome?

 Acrosome contains glycoprotein-digesting
enzyme, hyaluronidase & protease – helps in
penetration of protective layers surrounding egg
1
HMM/SCM 1424,CFS,IIUM
Development
Structure of spermatozoa
Egg Cell:
 Released as secondary oocyte at ovulation
 Surrounded by granulosa cells (forms the
corona radiata )
 Between granulosa cells & egg’s plasma
membrane is a glycoprotein (jelly) layer, zona
pellucida
Granulosa cells
(Corona radiata)
Zona pellucida
(jelly coat)
Egg membrane
Structure of egg cell
8.1.1 Stages of Fertilization

2
Development
Q: List four stages of fertilization.

Capacitation (iii) Cortical Reaction
Acrosome Reaction (iv) Activation
 In mammals, spermatozoa retained in

epididymis until their ejaculation
 Sperm head becomes covered with
glycoprotein coating → must be removed in
female reproductive tract before fertilization
can occur
(i) Capacitation
 Definition: Final biochemical event in the

maturation of mammalian spermatozoa. Required
to make spermatozoa competent to fertilize
oocyte.
 Non-mammalian- do not require this step,
ready to fertilize an oocyte.
 Enables spermatozoa to penetrate membranes
surrounding egg
 Stages of capacitation:
(a) removal of glycoprotein coat; &
(b) rearrangement of intramembranous
particles
 Characteristics of capacitated sperm:

High respiratory activity
(b) High motility
3
Development
(c) Ability to undergo acrosomal reaction
Acrosome Reaction
 Sperm must migrate through layer of follicle
cells/corona radiata- reaches the zona
pellucida.
 Head of the sperm contacts zona pellucida, the
acrosomal reaction is triggered .
 Steps within the acrosomal reaction:
(i) The acrosome discharges its contents by
exocytosis.
(ii) Hydrolytic enzymes digest zona pellucida
enable the acrosomal process to penetrate
the egg’s jelly coat.
(iii) Membrane protein of sperm binds to receptor
on egg membrane, membrane of sperm fuse
with membrane of egg.
 This lock-and-key recognition ensures
that eggs will be fertilized only by sperm
of the same species.
(iv) The entire sperm, tail and all, enters the egg.
(v) Enzymes harden zona pellucida
(iii) Cortical Reaction (Refer figure 47.6, pg 991)
4
Development
 Fusion of egg and sperm plasma membranes o

triggers release of Ca2+ from the egg’s ER into
cytosol.
 High concentrations of Ca2+ cause cortical
granules to fuse with the plasma membrane
and release their contents into zona pellucida
 Enzymes from granules thicken and harden the
zona pellucida. This resists entry of additional
sperms (slow block to polyspermy)
Q: What initiates the process of cortical

reaction?
The fusion of egg and sperm plasma
membranes
Activation
 High concentrations of Ca2+ in the egg :
5
Development
 stimulate increase in
rates of cellular respiration and protein
synthesis
 Cytoplasmic pH
changes from acidic to alkaline, due to
extrusion of H+
 Metabolism of activated egg
increases, the sperm nucleus swells and
merges with the egg nucleus, forming diploid
zygote.
8.2 Cleavage (Refer Fig. 47.7, pg. 992)
 3 successive stages following fertilization:

cleavage, gastrulation, & organogenesis
Definition of the cleavage process:

 Cleavage = successive rapid mitotic cell
divisions following fertilization, produces a
multicellular embryo, the blastula.
 No increase in overall size of zygote.
 The first five to seven divisions form a cluster
of cells known as the morula.
 A fluid-filled cavity called the blastocoel forms
within the morula- becomes a hollow ball of cells
called the blastula.
6
Development
 The zygote is partitioned into many smaller

cells called blastomeres.
 Each blastomere contains different
cytoplasmic determinants.
 Cleavage pattern is influenced by quantity &
distribution of egg-yolk.
 Yolk retards/slow down speed of cleavage –
thus, different region of the embryo may divide at
different rate.
Factors which affect the pattern of cleavage are:

(i) Quantity of yolk
(ii) Distribution of yolk
 Types of eggs based on the quantity of yolk:
(a) Isolecithal egg – little yolk (e.g. sea
urchin)
(b) Mesolecithal egg – moderate amount
of yolk (e.g. frog)
(c) Telolecithal egg – lots of yolk
concentrated at one end (e.g. bird)
 Polarity of embryo is determined by yolk

gradient:
(a) Vegetal pole – end with
highest concentration of yolk
(b) Animal pole – other end with
lowest concentration of yolk; represents
the most anterior part of embryo
7
Development
Different types of yolk distribution in

egg/embryo
 Cleavage patterns:
(a) Holoblastic
 Cleavage furrow extend completely
through egg
 Occur in isolecithal (sea urchin/human)
& mesolecithal egg (frog)
First 2 cleavage planes cut pole Then through meridional planes

to pole
8
Development
(i) (ii)
Holoblastic cleavage
(b) Meroblastic
 Cleavage incomplete & does not cut

through egg
 Occur in telolecithal egg (bird embryo)
Meroblastic cleavage
8.2.1 Comparisons of cleavage between

vertebrates
9
Development
A Primitive Chordates (Refer figure

47.7, pg 992)
 Little/no yolk (isolecithal egg)
 Holoblastic cleavage
 Symmetrical blastula with equally sized cells
(blastomeres)
 Example: lancets & agnathans
B Amphibians & Advanced Fish

 More yolk in vegetal hemisphere/pole
 Division of yolk-rich cells slow
 Holoblastic cleavage - assymetrical blastula
10
Development
 Large yolk-rich cells/blastomeres at vegetal pole

= macromere
 Small yolk-poor cells/blastomeres at animal pole
= micromere
 In frogs, the first two cleavages are vertical
and result in four blastomeres of equal size.
 The third division is horizontal, producing an
eight-celled embryo with two tiers of four cells.
(Refer figure 47.9,pg 993)
 The unequal distribution of yolk displaces the
mitotic apparatus and cytokinesis toward the
animal end of the dividing cells in equatorial
divisions.
 As a result, blastomeres of the animal
hemisphere are smaller than those in the
vegetal hemisphere.
 Continued cleavage produces a morula and
then a blastula.
 Because of unequal cell division, the
blastocoel is located in the animal
hemisphere.
Q: Why does cleavage produce blastomeres of

unequal sizes in frogs?
Due to unequal yolk distribution/ division of
yolk rich cells are slower/division of yolk poor
cells are faster
11
Development
Q: Label the figure below to indicate the animal

and vegetal poles of the frog’s embryo.
C Birds (Refer figure 47.10, pg 994)
 Egg composed entirely of

yolk, small amount of cytoplasm at animal
pole.
 Meroblastic cleavage
occurs at blastodisc (germinal disk) - a small,
circular, plate of embryonic cells/cytoplasm on
surface of yolk; it is where sperm enters egg;
nucleus of egg located here
12
Development
 Cleavage produces a cap of cells called the

blastoderm
 The blastomeres of the blastoderm sort into
upper and lower layers, the epiblast and the
hypoblast.
 The cavity between these two layers is the
blastocoel. (Blastula stage)
D Mammals
 Very little yolk, holoblastic cleavage – forms a
ball of cells, the trophoblast, surrounding a
blastocoel.
 Inner cell mass formed at one pole
 Mamalian blastula = blastocyst
13
Development
8.3 Gastrulation
 Gastrulation rearranges blastula to form a

three-layered embryo with a primitive gut/
archenteron = gastrula
 Involves movement of cells from periphery to
the inside of embryo
 Cells invaginate (dent inwards) & involute (roll
inward) into blastocoel.
 3 primary germ layers formed at end of
gastrulation: ectoderm, mesoderm, endoderm
Q: State the fate of each germ layer.
(Refer Figure 47.16: Fate of germs layers)

Germ layers Fate of layers
Ectoderm Epidermis, CNS, sense organs,
(Outer) neural crest
Mesoderm Dermis, skeleton, muscles, blood
14
Development
(middle) vessels, heart, gonads, kidneys

Endoderm Lining of digestive & respiratory
(Inner) tracts, liver, pancreas
 Endoderm surrounds the primitive gut

 Movement of cells brought about by contraction
of actin filaments
 Cell surface molecules (polysaccharides)
enable cells to stick to one another & move as
a single mass
 Gastrulation depends on shape of blastula
produced during cleavage
8.3.1 Gastrulation pattern in various animals
Primitive Chordates (Refer

Fig. 47.11, pg 995)
 Symmetrical blastula
 Invagination forms a 2-layered cup-shaped
gastrula
 Outer layer – ectoderm, Inner layer – endoderm
The stages of gastrulation of sea

urchin/primitive chordate.
(a) Begins with the migration of mesenchyme
cells from the vegetal pole into blastocoel.
(b) Vegetal pole invaginates. Mesenchyme cells
migrates through blastocoel.
15
Development
(c) Endoderm form from archenteron. New

mesenchyme at the tip of the tube begin to
send out thin extensions (filopodia) toward
the ectoderm of blastocoel.
(d) Contraction of filopodia drags the archentron
across blastocoel
filapodia extended by the secondary mesenchyme cells located at the tip

of the archenteron. This high magnification view shows a filopodium
extended by a secondary mesenchyme cell.
(e) Archenteron fuse with blastocoel to form

digestive tube with a mouth and anus.The
open end of archenteron, the blastopore, will
become the anus,a second opening forms
when the other end forms the mouth.
invagination
Gastrulation in primitive chordates
Neural plate
16
Future
Future
notochord
mesoderm
Archenteron
Endoderm
Ectoderm
Development
(2) Frog /Aquatic Vertebrates

(Refer figure 47.12, pg 996)
 Assymetrical blastula
Q: Discuss the gastrulation process in the frog
 Begins with formation of crescent shaped

blastopore
 Cells on the dorsal
surface of blastopore roll over the edge of the
dorsal lip and into the interior of the embryo =
involution. (occur at both side of embryo)
 A 2-layered gastrula is
formed
 Archenteron replaces
blastocoel
 Mesoderm then
pushes between ectoderm & endoderm
17
Development
 Lip of blastopore is
filled with yolk cells, the yolk plug
Reptiles, Birds, & Mammals

(Refer Figure 47.13, pg 997)
Avian/bird development process from

cleavage to gastrulation. (Refer Figure
47.17, pg. 999)
 Cleavage produces blastoderm atop mass of

yolk (Refer Figure 47.10, pg 995)
 Blastoderm differentiates into:
18
Development
 epiblast (upper layer)

 & hypoblast (lower layer)
 Cavity between = blastocoel
 During gastrulation, (Refer Figure 47.13, pg.

997) cells of epiblast migrate inward through
the primitive streak at midline
 Primitive streak is analogous to blastopore
 Some cells move laterally →mesoderm
 Others migrate downwards → endoderm
(displaces hypoblast)
 Epiblast that remain on top surface→ectoderm
Q: What happens to the hypoblast during post

gastrulation?
 Segregates/move away from endoderm
 Becomes part of the sac/membrane that
surrounds yolk and part of
stalk/connection between yolk mass and
embryo.
8.4 Neurulation/ Early organogenesis (Refer
figure 47.14, pg 997)
 Neurulation: Formation of embryonic neural

plate & its transformation into neural tube
 Embryo at this stage = neurula
 Differentiation of germ layers into tissues &
organs = organogenesis
19
Development
 Organogenesis involves morphogenetic

changes in tissue and cell shape
A Formation of notochord and neural plate

(Refer figure 47.14 (a), pg 997)
 Notochord formed from dorsal mesoderm that
condenses above archenteron
 Notochord - flexible rod (replaced by
vertebral column in vertebrates)
 Next to be formed is neural chord/neural tube
 Signals sent from the notochord to the
overlying ectoderm cause that region of
ectoderm to thicken and become neural plate.
 This process is often seen in
organogenesis: one germ layer signaling
another to determine the fate of the
second layer.
B Formation neural tube (Refer Figure 47.14

(b)
pg 997)
 Neural plate invaginates to form neural groove
 Edges of neural groove/ neural folds move
towards each other→ fuse →forming long hollow
neural tube (located beneath the surface of
embryo’s back)
20
Development
 Tube differentiates into brain/spinal cord/

spinal nerves
 In vertebrates, just before neural groove
closes, some cells from dorsal margin of neural
tube differentiate into neural crest (Figure 47.14
(b), pg. 997)
 Neural crest develops into gill arches, sensory
ganglia, Schwann cells, peripheral nerves,
teeth, skull, bones, & adrenal medulla
Neural
Neural foldplate
(A (C
) )
(B
)
Q: Label the figure above (Refer figure 47.14, pg

997)
(C) Formation of somites (Refer figure 74.14 (c)
pg 997)
Q: What are somites and where are they located?
 Somites are strips of mesoderm located next
to/lateral to the notochord.
 Are arranged serially on both sides along the
length of the notochord.
21
Development
 Mesenchyme cells migrate from the somites to

new locations.
 The notochord is the core around which the
vertebrae form.
 Parts of the notochord persist into adulthood
as the inner portions of vertebral disks.
 Somite cells also form the muscles associated
with the axial skeleton.
 Lateral to the somites, the mesoderm splits
into two layers that form the lining of the
coelom.
8.6 Extra-Embryonic Membranes (Refer figure

47.17 , pg 999)
 Consists of :
 yolk sac
 amnion
 chorion
 allantois
 In eggs of reptiles, birds, & uterus of mammals

or eggs/uterus of amniotes
 Embryos are surrounded by fluid within amnion
 Extra embryonic membranes : Not part of
developing animal – formed from tissues that
grow out from embryo
22
Development
(1) Yolk Sac

 Endoderm & inner mesoderm layer
(splanchnic) expand towards vegetal pole of
yolk & completely surround it, forming yolk sac
(bilaminar membrane)
 Function? Stores nutrients – transported to
embryo via blood vessels in yolk sac
membrane
(2) Amnion and Chorion

 Ectoderm & outer mesoderm layer (somatic)
grows over embryo & fuses above it
 The layers separate at point of fusion, producing
2 bilaminar layers:
(a) Amnion - inner membrane

 Encloses embryo in fluid-filled amniotic
cavity
 Function? Protects embryo from drying out
(b) Chorion - outer membrane

 Completely surrounds embryo & yolk sac
when cavity (extra-embryonic coelom)
between yolk sac & chorion expands
toward vegetal pole
 Next to eggshell
23
Development
 Function? Protects (with amnion) embryo

against mechanical shock
(3) Allantois
 Develops as evagination of embryo’s hindgut
 Consists of endoderm (inner layer) & mesoderm
(outer layer)
 Allantois expands within cavity between
chorion & yolk sac
 External surface eventually fuses with the
chorion forming a 3-layered chorioallantoic
membrane
 Functions?
1. Disposal sac for embryo’s uric acid (bird)
2. Forms respiratory organ for embryo –
embryonic blood vessels in allantois are
brought close to porous eggshell for gas
exchange(bird & reptiles)
3. Point of connection between embryo and
amnion, yolk sac, & allantois becomes
constricted as a vascularized umbilical cord
(mammals)
24
Development
8.7 Human Development

8.7.1 Mammalian/Human Early Embryonic
Development (Refer figure 47.18,pg. 1000)
 Fertilization in oviduct
 Cleavage occurs as embryo migrates to uterus
through oviduct
Q : State the major stages of early development of

humans.
Blastocyst reaches uterus
25
Development
()2 Blastocyst implants

()3 Gastrulation and formation of
extraembroynoic membrane.
Q: Identify 2 processes which help the embryo to

move through the oviduct/fallopian tube
towards the uterus. (Refer figure 46.15, pg.
979)
1 peristaltic contractions of the
oviduct
2 sweeping cilliary
motions/movements
Q: Describe the episodes of early cell

division/cleavage for human:
 Cleavage starts at 24 hours after fertilization
Division Completion time (hours Cell number
post fertilization) stage
1st 36 2 cells
2nd 60 4 cells
3rd 72 8 cells
 Cleavage is holoblastic, producing blastomeres
of equal sizes
 Morula/ball of cells reach uterus 3-4 days after
fertilization
 Cleavage continues, morula floats in uterus for a
few days
26
Development
 Blastocyst (blastula) is formed 7 days after

fertilization
Q: Describe the structures within a human

blastocyst.
1 a sphere of cell/ball of cells with a cavity (>
100 cells)
2 cavity is called blastocoel
3 contains Inner cell mass (ICM) - formed at
one end of blastocyst
4 a layer of cell - trophoblast surrounds
the blastocoel/cavity
Implantation and gastrulation (Refer notes for

Figure 47.18, pg. 1000)
 blastocyst starts to become implanted into the

endometrium ( 7 days post fertilization)
27
Development
 2nd week onwards/ 8 days onwards:

Trophoblast implants into endometrium & becomes
chorionic membrane
 ICM rearranges to form an amniotic cavity and
a disc of cells (below cavity)
 Disc rearranged to form epiblast (upper) and
hypoblast (lower)
 Gastrulation begins when cells from epiblast
invaginates inwards through primitive streak
forming mesoderm and endoderm
 Extraembryonic membranes also start to form
Trophoblast + mesoderm + adjacent
endometrium → placenta
 Hypoblast → yolk sac
Q: Can we assume that the human embryo

mainly originates from the epiblast?
Yes!
28
Development
The human extraembryonic membranes:

(1) Chorion
 Formed from trophoblast & mesoderm
 Surrounds embryo and all extra-embryonic
membranes
 Chorion forms finger-like chorionic villi –
project into blood spaces in uterine wall
 Chorion becomes chorionic frondosum (fetal
placenta)
 Surrounding epithelial tissues becomes decidua
basalis (maternal placenta)
 Chorionic fondosum + decidua basalis = placenta
 Mother’s & embryo blood close together in
placenta but don’t mix
Chorionic
frondosum
Placenta
Decidua
basalis
The human placenta
29
Development
Q: List out the functions of the placenta.

(1) Gaseous exchange – O2 from mother to
embryo; CO2 in opposite direction
(2) Provides nourishment for embryo
(3) Detoxify certain molecules that pass into
embryo
(4) Secretes hormones: HCG
HCG : Maintain corpus luteum → corpus
luteum secrete estrogen & progesterone
→ prevent menstruation & ovulation
Amnion
 Innermost membrane
 Formed from ectoderm & mesoderm
 Formed as a dome above epiblast
 Lines the fluid-filled amniotic cavity
 Encloses & cushions embryo in a fluid-filled
cavity (Fluid = amniotic fluid)
Yolk sac
 Formed from mesoderm & endoderm
 Encloses a fluid-filled cavity but no yolk
 Site for the formation of early blood cells
Diminishes as placenta develops & expands
Allantois
 Very small
30
Development
 Formed from mesoderm & endoderm

 Develops from outpocketing of gut
 Fuses with chorionic villi
 Incorporated into umbilical cord as embryo
grows
 Forms blood vessels – transfer oxygen and
nutrients from placenta to embryo & waste
products from embryo to placenta
31
Development
8.7.2 Complete Human Development
 Developing offspring:
 1st 8 weeks/1st 2 months = embryo
 Last 7 months = fetus
 Development/gestation (≈ 266 days) divided into
3 trimesters of ~ 3 months each
A. First Trimester
The First Month
1st week:
st
 1 cleavage of zygote – 30 hours after
fertilization
nd
 2 cleavage – 30 hours later
th th
 At implantation (7 – 10 day), embryo is called
blastocyst, with 128 to 256 cells
 Amnion & chorion formed
2nd week:
 Chorion forms chorionic frondosum while
surrounding epithelial tissues form decidua
basalis
 Gastrulation – primitive streak forms;
differentiation of germ layers
3rd week:
 Neurulation
32
Development
 First somites appear – develop into muscles,

vertebrae, & connective tissues
 Somites = blocks of mesoderm lateral to
notochord
 12 dozen somites at end of week – development
of blood vessels & gut
 Embryo ≈ 2 mm long
4th week:
 Embryo obtains nutrients directly from
endometrium
 Organogenesis – formation of eyes & 4-
chambered heart
 Heart starts to beat
 Over 30 pairs of somites available
 Arm & leg buds form
 Embryo ≈ 5 mm long
33
Development
34
Development
Second Month
 Morphogenesis – formation of shape

 Limbs assume adult shape
 Bones of embryonic tail fuse to form coccyx
 Major organs evident in abdominal cavity
including liver, pancreas, & gallbladder
 Embryo ≈ 25 mm long, 1 g
 Looks distinctly human
35
Development
Third Month
 Nervous system & sense organ develop
 Arms & legs move
 Facial expressions
 Primitive reflexes – startle reflex & suckling
 All major organs established → growth
 At 10th week, HCG secretion by placenta
declines – corpus luteum degenerates
 But placenta secretes estrogen &
progesterone:
(i) Inhibits menstruation
(ii) Inhibits FSH & LH – prevents ovulation
(iii) Maintains uterus & prepare it for labor &
delivery
(iv) Stimulate development of
mammary glands – preparation for
lactation
36
Development
Third months
2. Second Trimester
 Rapid growth & development
 Bones enlarge
 Baby starts kicking
 Head & body covered by fine hair (lanugo )
which is then lost
 Rapid heart beat at end of 5th month
 Fetus ≈ 175 mm long, 225g
 End of 6th month ≈ 30 cm long, 600 g
37
Development
4th months
5th months
38
Development
6th months
3. Third Trimester
 Period of growth
 Body weight increases 500 – 600%
 Brain & peripheral nervous system enlarge –
mature rapidly
 Fetus regulates own temperature & controls
own breathing
 Fat deposits form under skin giving rounded,
chubby shape characteristic of many babies
 In male, testes descend into scrotum
 Immune system does not mature by time of
birth
39
Development
 During final fetal month, antibodies from

mother’s blood cross placenta – provides
temporary defenses against infectious diseases
 Breast-fed babies receive maternal antibodies
& immune cells in breast milk
 At birth weight of baby maybe 3-4 kg, and ~ 50
cm in length
7th months
40
Development
8th months
9 months
41
Development
Critical Periods of Human Development
Image Source: http://www.people.virginia.edu/~rjh9u/hdevsum.html
8.8 Parturition and lactation

8.8.1 Parturition
 Parturition – the act of giving birth

Q: State the three stages of labor
Three main stages: (Refer Figure 46.19, pg.,
981)
42
Development
(1) First Stage: Dilation of the cervix
≈ 12 hours (8 – 24 hours for 1st pregnancy, but

may be a few minutes after many
pregnancies); cervix thins and expands/dilates
– baby’s head passes into vagina; amniotic
fluid released from uterus.
(2) Second stage: Expulsion/Delivery of infant

≈ 20 – 60 minutes; uterine contraction
stronger & more frequent; baby is pushed
through vagina & out of mother’s body.
43
Development
(3) Third stage: Delivery of the

placenta/Afterbirth –
≈ 10 – 15 minutes; further uterine contractions
push umbilical cord & placenta out of mother’s
body.
44
Development
Process of parturition
 Initiation of birth:
(1) Fetal pituitary gland secretes
adrenocorticotropic hormone (ACTH)
(2) Stimulates fetal adrenal glands to secrete
coticosteroids
(3) Passes into maternal placenta→ stimulates
estrogen production & inhibits progesterone
production.
(4) Changes in estrogen & progesterone levels
stimulates production of prostaglandin by
placenta. Estrogen level normally reaches
highest level (in mother’s blood) during last
weeks of pregnancy
(5) Estrogen also induces formation of oxytocin
receptors on uterus
(6) Estrogen also stimulates release of oxytocin
by maternal posterior pituitary gland
(7) Oxytocin causes muscles of uterus to
contract
(8) At the same time, fetal head pushes against
cervix
(9) Stimulates stretch receptors
(10) Send information to mother’s brain →
stimulates release of more oxytocin from
posterior pituitary gland
(11) Causes uterine muscle to contract more and
stimulates placenta to secrete prostaglandins
45
Development
(12) Prostaglandins further enhance uterine

contractions
(13) Levels of prostaglandins & oxytocin increase
during labor due to the stimulation of physical
& emotional stress during labor ( positive
feedback system)
(14) Frequency & intensity of contraction
increases gradually, leading to stronger and
rhythmic contractions
(15) Finally, contraction pushes baby out of
uterus.
46
Development
8.8.2 Lactation
 Secretion of milk from mammary gland

Q: Name two hormones involved in lactation.
 2 hormones involved:
(i) Prolactin – promotes milk production
(ii) Oxytocin – stimulates ejection of milk
from nipples
 Prolactin secreted by anterior pituitary gland
 Its effect inhibited by progesterone & estrogen
 After birth, levels of progesterone & estrogen fall
 Enables prolactin to promote milk production
 Its level increases temporarily every time baby
suckles from mother
 Thus, milk supply is regulated to meet demands
of baby
 The more a baby feeds, the greater the prolactin
secretion & milk production – positive
feedback
 Suckling also:
(i) Triggers release of oxytocin – ensures milk
flow freely
(ii) Inhibits secretion of LH & FSH – process is
called lactational anovulation → Suppresses
follicular development & ovulation
 Milk production stops after a mother stops
breast-feeding.
8.9 Plant Development
47
Development
8.9.1 Seed Development
Endosperm Development (Refer pg. 777)
 Develops before zygote/embryo.

 Triploid nucleus divides repeatedly, forming
multinucleate cell/supercell of milky consistency
 Membranes then form between nuclei
(cytokinesis), forming multicellular endosperm.
 Endosperm becomes solid
 Contains nutrient for developing embryo:
 Starch – produced by endosperm.
 Lipid & protein – passed to endosperm by
parent plant.
 In most monocots, nutrients used by embryo
during germination.
 In most dicots/eudicots, nutrients exported to
cotyledons – thus, mature seeds lack
endosperm.
48
Development
Embryo Development (Refer Fig. 38.7, pg. 777)
 First division is transverse & asymmetric,

forming smaller terminal cell & larger basal
cell.
 Basal cell continues dividing transversely
forming a row of cells called suspensor.
 Suspensor transfers nutrients from parent plant
or endosperm to embryo.
 Terminal cell divides repeatedly forming a
globular/spherical proembryo attached to
suspensor.
 Early formation of cotyledons in dicots gives
embryo a heart-shape appearance, and finally
as embryo elongates, it changes into a
torpedo-shaped embryo.
 Embryo differentiates into 3 main parts:
 radicle (young root),
 plumule (young shoot),
 1 or 2 cotyledons (young leaves)
.
49
Development
Structure of a mature seed

 Seed dehydrates at late maturation. 5-15% of its’
weight is water
 Embryo surrounded by food supply
(cotyledone/endosperms/both)
 Embryo becomes dormant
Q: Define dormancy in seeds.

A condition whereby growth and metabolism
ceases/ stops
 Embryo + food = enclosed by seed coat

(originated from integument of ovary)
 Ovule develops into seed
 Hypocotyl = below attachment of cotyledons
 Radicle = terminal end of hypocotyl→embryonic
root
 Epicotyl = above attachment of cotyledons,
contains shoot tip, with a pair of miniature
leaves
 Members of grass family (monocot) have
specialized cotyledon = scutellum
 Embryo of grass contains 2 coverings:
 Coleoptile : covers young shoot
 Coleorhiza: covers young root
From ovary to fruit

 Ovary → flesh of fruit
50
Development
 Ovary wall → pericarp= thickened wall of fruit

 A fruit ripens at the same time as seeds within it
complete their development
8.9.2 Seed Germination
 Dormancy increases chances that germination

will occur at a time & place most
advantageous to seedling.
 Breaking dormancy & stimulation of germination
requires certain environmental conditions –
water, suitable temperatures, light, & growth
promoters.
 Duration of dormancy varies.
Germination
Example : germination of cereal grains
(a) Imbibition occurs, seed

takes in water rapidly
(b) Cellular respiration in seed
starts
(c) Embryo secretes gibberellin
hormone /GA into endosperm
(d) GA acts on aleurone layer,
stimulating aleurone to synthesize and
secrete digestive enzymes
51
Development
(e) Enzymes digest/hydrolyze

stored food in endosperm into soluble
molecules/monomers.
amylase
Starch → disaccharides/sugar
Proteinase/peptidase
Protein → peptides/amino acids
lipase
Lipid → fatty acid + glycerol
(f) nutrients mobilized by the

enzymes are absorbed by scutellum
(g) nutrients transferred to
embryo
(h) nutrients are used by
embryo for energy/ growth/ synthesis of
polymers/ – translocated to plumule &
radicle for germination
Endosperm
Cotyledon
Hydrolysis
Plumule Sugar Starch
α -amylase
Translation
RNA
Transcription
DNA
Amino acids
52 Hydrolysis
Radicle HMM/SCM 1424,CFS,IIUM
Storage protein
Imbibition
H2O
Development
8.9.3 Growth of Seedlings
 Germinating seed continues to undergo

metabolic changes culminating into a seedling
 Two types of seedling growth:
(i) Epigeal germination
 Example: sunflower, long beans,(many
eudicots/dicots)
 Elongation of hypocotyl (area below
cotyledon) pushes epicotyl and
cotyledon above soil surface
(Refer Figure 38.10(a), pg 780)
(ii) Hypogeal germination

53
Development
 Example: broad beans, maize, grass

(many monocots)
 Elongation of epicotyl (area above
cotyledon)/coleoptile pushes
plumule/shoot tip above soil surface
 Cotyledons remain underground
54
Development
(Refer Figure 38.10(b), pg 780)
8.10 Growth
8.10.1 Definition of Growth
 Growth is closely linked to development:

 Growth: permanent & irreversible increase
in dry mass of living material
 Development: increase in complexity
 3 phases of growth:
(i) Cell division (hyperplasia ) – increase in
cell number
(ii) Cell expansion (hypertrophy ) –
irreversible increase in cell size as a
result of uptake of water or synthesis of
living material
(i) Cell differentiation – specialization of
cells
 All phases involve biochemical activity,

especially protein synthesis
 Growth & development determines the process
of morphogenesis – changes in overall form &
structure of individual organs & of the
organism as a whole
55
Development
 Positive growth: anabolism > catabolism

Increase in human height

 Negative growth: catabolism > anabolism
 Example: germination
 Positive (increase) – cell size, length, &
volume
 Negative (decrease) – dry mass
8.10.2 Measuring Growth
 Measured at different levels – community to

molecular levels
 Parameters at level of organism – length, area,
volume, surface area, number, & mass (fresh
or dry)
 Fresh mass:
 Advantage:
(a) Easier to measure – requires
less preparation of sample
(b) Doesn’t injure organism –
repeated measurement can be made
 Disadvantage:
Data inconsistent – fluctuations in water
content
 Dry mass:
 Organism is killed - placed in oven (110°C),
cooled, & weighed
 Procedure repeated until constant value is
recorded
56
Development
 Difficult to carry but more accurate data
Q: What are the advantages and disadvantages of

using dry mass measurements to determine
growth?
 Advantage:
Accurate measurement of the amount of
organ matter present.
 Disadvantage:
(a) Organisms are killed, growth of
same specimen cannot be measured
continuously
(b) Needs a large number of
specimens to measure growth change
(c) Time consuming (preparation of
sample)
57
Development
8.10.3 Typical Growth Curve
 Sigmoid curve
Growth I II III
parameter IV
Inflexion point
Time
The Sigmoid Curve
 4 phases:
(I) Lag phase
 Slow positive growth as organisms
acclimatize to the environment or it is a
period of preparation of growth
(II) Logarithmic (Exponential) phase

 Rapid population growth
 Maximum growth rate/no constraint on
growth
 Adequate nutrients and no accumulation of
waste products
58
Development
 Point at which exponential growth begins

to decrease = inflexion point
(III) Decelerating phase

 Growth rate slows down due to internal or
external factors or interaction of both
(IV) Plateau/Stationary phase

 Dynamic equilibrium or steady state
 Parameter fluctuates but tend to be equal
over time
 A period leading to senescence and death
Nature of curve varies:
(a) May increase slightly until organisms die

 Example: monocot leaves, non-vertebrates,
fish, & certain reptiles
 Positive growth
(b) May flatten – no change in growth

 Example: certain Cnidarians
(c) May tail off/decrease

 Example: many mammals
 Negative growth – leads to senescence
59
Development
8.10.4 Types of Growth Curve
Example: Growth measurements by height of plant
Days Height/cm Growth rate Relative

after (increase in growth rate
planting height (increase in
during each height as % of
10-day growth at time
period) of
measurement)
10 2 2 100
20 7 5 71
30 20 13 65
40 40 20 50
50 75 35 46
60 110 35 31
70 140 30 21
80 150 10 7
90 155 5 3
100 160 5 3
110 160 0 0
120 160 0 0
1 2 3 4
60
Development
(1) Actual/Absolute Growth Curve
 Definition:Parameter against time

 Shows overall growth pattern & extent of growth
Height/cm
Days
The absolute growth curve
(2) Absolute Growth Rate Curve

 Definition: Changes in parameter against time
– increase in parameter that take place during
successive intervals of time are plotted against
time
 Rate of growth changes with time
 Rate increases steadily, reaches a maximum,
& gradually falls
 Steepest part of curve – period of most rapid
growth
 Peak = inflexion point – after this point rate of
growth decreases
61
Development
Increase in height
during 10-day
period/cm
Days
The absolute growth rate curve
(3) Relative/Specific Rate of Growth

 Definition: Increase in growth is expressed as
a percentage of the previous amount
 Value is then plotted against time
Gain as % of
previous height
Days
The relative growth rate curve
62
Development
 Growth is fastest in the beginning of life, after

which it gradually slows down
Example:
 Between ages 1 – 2 years, a baby’s mass might
increase from 10 to 12 kg
 Absolute increase = 12 – 10 = 2 kg
 Relative increase = (2/10) x 100 = 20%
 In the same period of time, mass of teenager
increases from 50 to 55 kg
 Absolute increase = 55 – 50 = 5 kg
 Relative increase = (5/50) x 100 = 10%
Growth in Mammals
 Absolute growth curve – sigmoid

 4 distinct phase of human growth – embryo,
infancy/childhood, adolescence, & adulthood
63
Development
Embryo Infancy/childhood Adolescence Adulthood
Greatest growth rate-

70
infancy (first 2 years) &
Mass/kg Adolescent
(m) Fetal adolescent (13-17 years) growth
50 growth
30 Juvenile growth
10 Infant growth
0 2 4 6 8 10 12 14 16 18
20 22
Birth Age/years (t)
Conception
Change in
Maximum mass –adulthood
mass/kg 6 Greatest change of mass, 0-2 years
(dm/dt) & 9- 14 years
4
0 0 2 4 6 8 10 12 14 16 18
20 22
Age/years (t)
Greatest relative growth rate –

Relative embryological development
change in
1.0
mass/kg
(dm/dt.1/m)
0.5
0 2 4 6 8 10 12 14 16 18
20 22
Age/years (t)
Human growth curve
64
Development
Patterns of Growth
1. Isometric and Allometric growth
(a) Isometric Growth

 “isos” = same; “metron” = measure
 Organs grow at same mean rate as rest of
body
 Change in size of organism not accompanied
by change in shape or external form of organism
 Proportion of structures remain constant
 Example: salamander, fish, & some insects
 Little change in shape but marked change in
mass
 Example: length increases by 10% while mass
increases by 33%
Isometric growth in a salamander
(b) Allometric Growth

65
Development
 “allos” = other
 Organs grow at different rate from rest of body
 Rapid growth of lymph tissues during
childhood – provides young organism with
natural immunity & defense against disease
 Last organ to develop & differentiate are the
reproductive organs
Allometric growth as shown by human organs and

tissues
 Change in size of organism accompanied by

change in shape of organism
 Example: mammal
66
Development
Allometric growth in human male
2. Limited and Unlimited Growth
(a) Limited
(Definite/Determinate) Growth
 Period of maximum growth (maturation &

reproduction) followed by period of negative
growth (senescence) before death
 Example: annual plants, insects, birds, &
mammals
Dry mass/g
Senescence
(Negative growth)
Time/weeks
67
Development
Limited growth
(b) Unlimited
(Indefinite/Indeterminate) Growth
 Growth curve shows a cumulative series of

sigmoid curves
 Each curve represents one year’s growth
 Growth continues until death
 Example: fungi, algae, non-vertebrates,
fishes, reptiles, monocot leaves
Height/m
Time/years
Indefinite growth
8.10.3 Growth in Arthropoda
 Growth occurs in spurts interrupted by a series

of moults – discontinuous growth
 Due to inelastic nature of exoskeleton
 Prevents body from growing
 Exoskeleton is shed periodically through
moulting (ecdysis)
68
Development
Adult
Length/cm 5th
instar
4th
3rd instar
2nd instar
1st
instar
instar
Time/days
Discontinuous growth
69

Animal Development

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Development

Q: What is the importance of acrosome?

Structure of egg cell

8.1.1 Stages of Fertilization

Q: List four stages of fertilization.

 In mammals, spermatozoa retained in

 Definition: Final biochemical event in the

 Characteristics of capacitated sperm:

(c) Ability to undergo acrosomal reaction

(iii) Cortical Reaction (Refer figure 47.6, pg 991)

 Fusion of egg and sperm plasma membranes o

Q: What initiates the process of cortical

8.2 Cleavage (Refer Fig. 47.7, pg. 992)

 3 successive stages following fertilization:

Definition of the cleavage process:

 The zygote is partitioned into many smaller

Factors which affect the pattern of cleavage are:

 Polarity of embryo is determined by yolk

Different types of yolk distribution in

First 2 cleavage planes cut pole Then through meridional planes

 Cleavage incomplete & does not cut

8.2.1 Comparisons of cleavage between

A Primitive Chordates (Refer figure

B Amphibians & Advanced Fish

 Large yolk-rich cells/blastomeres at vegetal pole

Q: Why does cleavage produce blastomeres of

Q: Label the figure below to indicate the animal

C Birds (Refer figure 47.10, pg 994)

 Egg composed entirely of

 Cleavage produces a cap of cells called the

 Gastrulation rearranges blastula to form a

Q: State the fate of each germ layer.

(Refer Figure 47.16: Fate of germs layers)

(middle) vessels, heart, gonads, kidneys

 Endoderm surrounds the primitive gut

8.3.1 Gastrulation pattern in various animals

Primitive Chordates (Refer

The stages of gastrulation of sea

(c) Endoderm form from archenteron. New

filapodia extended by the secondary mesenchyme cells located at the tip

(e) Archenteron fuse with blastocoel to form

Gastrulation in primitive chordates

(2) Frog /Aquatic Vertebrates

Q: Discuss the gastrulation process in the frog

 Begins with formation of crescent shaped

Reptiles, Birds, & Mammals

Avian/bird development process from

 Cleavage produces blastoderm atop mass of

 epiblast (upper layer)

 During gastrulation, (Refer Figure 47.13, pg.

Q: What happens to the hypoblast during post

 Neurulation: Formation of embryonic neural

 Organogenesis involves morphogenetic

A Formation of notochord and neural plate

B Formation neural tube (Refer Figure 47.14

 Tube differentiates into brain/spinal cord/

Q: Label the figure above (Refer figure 47.14, pg

 Mesenchyme cells migrate from the somites to

8.6 Extra-Embryonic Membranes (Refer figure

 In eggs of reptiles, birds, & uterus of mammals

(1) Yolk Sac

(2) Amnion and Chorion

(a) Amnion - inner membrane