Unit – 2 Lecture – 1

GAMETOGENESIS
Course : B.Sc. (H) Zoology VI semester
Paper : DEVELOPMENTAL BIOLOGY

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 SYLLABUS

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 SYLLABUS

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 Questions to address…..

• What is gametogenesis?
• How does spermatogenesis take place?
• What are the various phases of oogenesis?
• What are the various types of eggs?
• What membranes do the eggs of different animals
bear?

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 GAMETOGENESIS
Spermatogenesis
Oogenesis

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 GAMETOGENESIS

• The process of formation of gametes is called gametogenesis.

• In the male, it is called spermatogenesis, takes place in the seminiferous
tubules of the testes and it begins at puberty.
• In the female, maturation from primitive germ cell to mature OVUM,
which is called oogenesis, begins before birth
• The mammalian Primordial Germ Cells (PGCs) are bipotential and can
become either sperm or eggs; depending upon whether they reside in the
ovaries or testes resp.
• All of these decisions are coordinated by factors produced by the
developing gonads.
Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 • PGCs do not originally form inside the gonads, rather they
originate in the posterior portion of the embryo and migrate
into the gonads via hindgut in the 5th week of development
• During this migration, the PGCs are surrounded by cells
secreting stem cell factor (SCF). These cells form a ‘travelling
niche’ around the PGCs. SCF is necessary for PGC motility and
survival.

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 Gametogenesis involves 3 phases:

• Multiplication/Proliferation phase
PGCs (spermatogonia/oogonia) multiply & proliferate by mitosis
• Growth phase
increase in cell size to become primary spermatocytes/oocytes
• Maturation phase
meiosis to produce mature gametes
in both spermatogenesis & oogenesis

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 SPERMATOGENESIS
THE PROCESS OF FORMATION OF MATURE MALE
GAMETE (SPERM)

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 SPERMATOGENESIS
Initiates in the embryonic testes:
• mammalian PGCs (then called GONOCYTES =true stem cells)
arrive at the genital ridge of a male embryo &and become
incorporated into the sex cords that will become the
seminiferous tubules
But these PGCs remain dormant during childhood and actively begin
producing sperm by meiosis at puberty.
Stra8 transcription factor, which promotes a new round of DNA
synthesis and meiotic initiation in the germ cells & Retinoic acid
produced by the mesonephros kidney are essential for initiating
meiosis.
Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 SPERMATOGENESIS

It is the process of formation of

mature sperms (spermatozoa) in
seminiferous tubules of testis. It
has 3major phases:
a. Proliferative phase where sperm
stem cells (spermatogonia)
increase by mitosis.
b. Meiotic phase, involving the two
divisions that create the haploid
state
c. Spermiogenesis or the
postmeiotic “shaping” phase
during which the round cells
(spermatids) transform
into spermatozoa while they go
through a series of changes

Course: Developmental Biology Created by:Figure

Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
6.22 Gilbert 11th edn.
 1.PROLIFERATIVE PHASE
• gonocytes ➔ become undifferentiated
Spermatogonia residing near the basal end of
the tubular cells
• Spermatogonia differentiate into Type A
(A1,A2,A3,A4) Spermatogonia, by mitotic
proliferation
• Spermatogonia keep attached to Sertoli cells or
Sustentacular cells
• Sertoli cells serve to nourish the developing
sperm
• In the spaces between adjacent seminiferous
tubules are clusters of cells called interstitial
cells / Leydig cells - secrete Testosterone
• Testicular blood vessels serve to vascularize the
process

Course: Developmental
Figure 6.22 Biology
Gilbert 11 th edn. Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 Sertoli cells or Sustentacular cells (in the epithelium of the
seminiferous tubules; extend from the basement membrane to the
Sperm maturation. (Figure 6.21 Gilbert 11th edn.) lumen of the tubule) by adhesion molecules.
(A) Cross section of the seminiferous tubule.
(B) Simplified diagram of a portion of the seminiferous tubule, illustrating relationships between spermatogonia,
spermatocytes, and sperm.
As these germ cells mature, they progress toward the lumen of the seminiferous tubule.
Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 2. The meiotic phase: Haploid spermatids
• Spermatogonia with high levels of Stra8 and
responding to Stem Cell Factor (SCF) divide
mitotically to become type B spermatogonia.
• Type B spermatogonia are the precursors of the
spermatocytes and are the last cells of the line
that undergo mitosis. They divide once to
generate the primary spermatocytes (2n) —the
cells that enter meiosis.
• Each primary spermatocyte undergoes the 1st
meiotic division to yield a pair of secondary
spermatocytes(n) that undergo the second
meiotic division.
• Haploid cells thus formed are called Spermatids
(n), and they are still connected to one another
through their cytoplasmic bridges. The spermatids
that are connected in this manner have haploid
nuclei but are functionally diploid, since a gene
product made in one cell can readily diffuse into
the cytoplasm of its neighbors
Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 • As the undifferentiated
spermatogonia divide to form
spermatids:
– they move farther and farther
away from the basal lamina of
the seminiferous tubule and
closer to its lumen
– they lose their cytoplasmic
connections and differentiate
into spermatozoa

Thus, Each Primary spermatocyte produces

2 secondary spermatocytes. Each secondary
spermatocyte produces 2 haploid
spermatids. Thus 4 spermatids are formed
from each primary spermatocyte.
Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 3. Spermiogenesis/Spermateliosis:
differentiation of the sperm
• For fertilization to occur, the sperm
has to meet and bind with an egg,
and spermiogenesis prepares the
sperm for these functions of
motility and interaction

Course: Developmental Biology11th edn.

Figure 6.22 Gilbert Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 Spermiogenesis…..

• Construction of the Acrosomal vesicle/Acrosome

from the Golgi apparatus: The acrosome forms a
cap that covers the sperm nucleus
• Nucleus flattens and condenses
• Nucleus rotates so that the acrosome points away
from the lumen of seminiferous tubule
• A centriole pair reaches the opposite end of
spermatid – gives rise to flagellum – which
extends into the lumen of seminiferous tubule
• Mitochondria aggregate to form a ring around the
base of the flagellum
• Histones of the spermatogonia are often replaced
by sperm-specific histone variants &
PROTAMINES (Arginine-rich proteins), and
widespread nucleosome dissociation takes place
• Excess cytoplasm is shed as residual body, except
for the thin sheath – Manchette surrounding the
mitochondria along the posterior region of
Modification of a germ cell to form a mammalian sperm.
nucleus.
Figure 7.1 Gilbert 11th edn.
Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 • Sperms continue to develop even after they leave the testes
(epididymal maturation)
• final differentiation in the reproductive tract of female:
oviducal secretions from the oviducts (capacitation)
• Thus, the full differentiation of the sperm take place in two
different organisms.

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 • In the mouse, development from stem cell to spermatozoon takes
34.5 days:
– the spermatogonial stages last 8 days, meiosis lasts 13 days, and
– spermiogenesis takes another 13.5 days.
• Human sperm development takes nearly 65 days.
• Each day, some 100 million sperm are made in each human testicle,
and each ejaculation releases 200 million sperm.
• Unused sperm are either resorbed or passed out of the body in
urine.
• During his lifetime, a human male can produce 1012 to 1013 sperm

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 Role of Sertoli/Sustentacular cells
• support and protect developing
spermatogenic cells
• nourish spermatocytes, spermatids, and
sperm
• phagocytize excess spermatid cytoplasm
• control movements of spermatogenic cells
& release of sperm into the lumen of the
seminiferous tubule
• produce fluid for sperm transport
• secrete the hormone inhibin
• regulate the effects of testosterone and
follicle-stimulating hormone (FSH)

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 Draw diagrams:
1. Process of spermatogenesis.
2. Simplified diagram of a portion of the seminiferous tubule,
illustrating relationships between spermatogonia,
spermatocytes, and sperm.
3. Modification of a germ cell to form a mammalian sperm.
4. Structure of human sperm.
5. Axial filament (T.S.) along with explanation.

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 STRUCTURE OF SPERM
• highly specialized cell
• greater part = nuclear material
• most cytoplasm = eliminated during maturation
• certain organelles left = modified for spermatic function
• propulsive system: Axial Filament (mitochondria)
• A plasma membrane envelops the whole body
• totally devoid of stored food and protective envelopes
• carry paternal chromosomal elements & activate ovum

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
Course: Developmental Biology Created
Structure of human by: Dr.Priya
sperm Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
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 • A typical mammalian spermatozoa consists of the
following main parts:
– Head
– Neck
– Middle piece
– Tail / flagellum

Human sperm, visualized using scanning electron microscopy,

have a flagellum, neck, and head Figure 43.9
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 Head -
spoon-shaped
3 main components: acrosome, nucleus, cytoplasm
• Acrosomal Vesicle or Acrosome:
– vesicles pinch off from golgi apparatus & coalesce to form acrosome
– Acrosomal Enzymes: Hydrolases, Hyaluronidase, Zona Lysin or Acrosin
• digest proteins and complex sugars
• lyse egg membranes
– @ fertilization = penetrate through egg membranes & egg activation

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Structure of human by: Dr.Priya
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 • Nucleus (n): During the course of sperm maturation
– streamlined
– DNA: tightly compressed
– major part of the head
– contains genes
– transmission of the
hereditary characters
from male parent
Structure of human sperm https://en.wikipedia.org/wiki/Sperm

• Cytoplasm forms a condensed layer known as the

MANCHETTE or Periacrosomal Cytoplasm around the
posterior part of the nucleus
Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 Neck -
• two CENTRIOLES (Proximal and Distal) at right angle to each
other
• Proximal Centriole → spindle system ➔ 1st cell division in the
fertilized egg
• Distal Centriole: → axial filament of the flagellum of the
spermatozoa → basal granule
• 9+0 pattern

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 Middle piece -
within outward :
– Axial Filaments / Axoneme: flagellar motor core = 9+2 pattern
– Mitochondrial sheath: spiral around the rings of filament; ATP
– Cytoplasm: Manchette
– Plasma Membrane
*Ring Centriole b/w middle piece & tail

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 Tail or Flagellum -
• within outward: Axial Filaments, Fibrous Sheath, Cytoplasm And
Cell Membrane
• Principal piece: fibrous helix surrounds the longitudinal fibres
• End piece: axial filaments only covered with cytoplasm and
plasmalemma

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Structure of human by: Dr.Priya
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 Functions of Sperm
• Contribute ½ genetic information to diploid offspring
• Restores diploidy in the offspring
• Mammals: sex of the offspring (XX-XY mechanism)
• Provides spindle for 1st cleavage of the zygote
• Activates the egg to complete its development

DIY-3
Sperm head is of various shapes & may be
flagellated or non-flagellated. Give examples with
diagrams.
Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 OOGENESIS
THE PROCESS OF FORMATION OF MATURE
FEMALE GAMETE (OVUM) IN THE OVARY

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 Figure 27.11 Oogenesis
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Figure 19.29
The ovarian follicle of mammals. (A) Maturation of the ovarian follicle. When mature, it is often called a Graafian follicle.

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 Summary of Oogenesis and Follicular Development

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 OOGENESIS: 3 PHASES:

1. MULTIPLICATION /PROLIFERATION PHASE: Before birth

• PGCs → mitosis → undifferentiated GCs OOGONIA (2n) by 3rd
month of gestation
• grow in size → PRIMARY OOCYTES by 7th month of gestation
• arrested in Diplotene of prophase-I of meiosis (=DICTYATE STAGE)
• → primordial follicle (surrounded by a layer of follicular cells)

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 2. GROWTH PHASE: till puberty
• Growth in primary oocyte: 100X
• Nucleus enlarges = Germinal Vesicle (GV)
• 2 phases: Previtellogenesis & Vitellogenesis
• Granulosa cells surround the primary oocyte
= Primary Follicle
The number of germ cells in the
human ovary changes over the life
span
th
Figure 6.23 Gilbert 11 edn.
Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 Antrum

• Follicular Fluid / Liquid Folliculi ➔

Follicular cavity/Antrum
• follicle is now called the Secondary /
Vesicular / Graafian follicle
Discus Proligerus

Cumulus Oophorus or Germ Hill

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
3. MATURATION PHASE:

• Luteinizing Hormone (LH) surge

• Chromosome condensation
• Germinal Vesicle Break Down
(GVBD)
• 15-20 follicles begin to mature in
each ovary alternate month ➔ one
reaches maturity → others ATRESIA
• Meiosis I completed →Secondary
oocyte
• BUT arrested at metaphase II
• Meiosis II completed upon
fertilization (High Calcium
inactivates cytostatic factor)
• Female pronucleus is formed Figure 27.11 Oogenesis
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Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 Graafian Follicle

Ovulation
Corpus Luteum
(theca lutein + granulosa lutein cells)

Fertilization No fertilization
Luteal cyst hyaline structure
Progesterone (implantation) Corpus Albicans
Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 GROWTH PHASE of primary oocyte is divided into two stages:
PREVITELLOGENESIS & VITELLOGENESIS

1. PREVITELLOGENESIS: no yolk formation, changes in cell

organelles
1.1 Changes in Nucleus
• Nucleus ↑ nucleoplasm ↑ = Germinal Vesicle
• Nucleoli ↑ Transcription of rRNA ↑ by Nucleolar Organizer
Region (NOR) of chromosomes
Informosomes (A.S.Spirin,1964): packed forms of
mRNA-protein complex = immediate production of
proteins by the zygote
Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 • Lampbrush chromosomes* (W. Flemming, 1882, Salamander
eggs)
– formed during diplotene of meiosis-I
– giant, looped, ↑ transcription of mRNA & tRNA ↑ protein
synthesis in the cytoplasm by gene amplification *(↑
selective gene expression)
– egg of some fishes, amphibia, reptiles & birds, absent in
sea urchins & mammals

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 1.2 Changes in CYTOPLASM
– Cytoplasm ↑
– Ribosomes ↑ protein synthesis ↑
– Mitochondria ↑
– ER / Annulate Lamellae ↑elaborate
• transfer of contents of nucleus into the cytoplasm
• disappear in late oogenesis

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
1.3 Cortical Granules: spherical, membrane bound vesicles
▪ formed from RER & Golgi
▪ in Sea Urchins, Fish, Frogs, Reptiles & some mammals
Cortical enzymes
✓ Acid Mucopolysaccharides
✓ Hyaline proteins : blastomere adhesion after cleavage
✓ Enzymes: Ovoperoxidase (OVOP – hardens the fertilization
membrane by crosslinking the Tyrosine residues & is
spermicidal)
▪ explode at the site of sperm entry, block polyspermy
▪ formed in the endoplasm, then move to the peripheral region
(cortex) of cytoplasm’
Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 1.4 Cortex : the peripheral cytoplasmic layer of the oocyte
– actin microfilaments = cell constriction during cleavage &
alter cell shape during gastrulation
– Microvilli = engulf sperm during fertilization
– semi-rigid & more viscous (cortical granules)
– Melanosomes (melanin) UV light filters

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 2. VITELLOGENESIS: the process of laying down yolk in the
primary oocyte (egg)
• diplotene stage of Meiosis-I
• Amount of yolk = ↓ mammalian egg; ↑↑ birds, reptiles &
shark egg
• Precursor of yolk = Vitellogenin (synthesised by mitochondrial
aggregates called Yolk nucleus of Balbiani as in oocytes of
spiders, amphibians, & birds)

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 Methods of vitellogenesis:
1. Autosynthesis: synthesis of vitellogenin by egg
2. Heterosynthesis: synthesis of vitellogenin by extraovarian
tissues (maternal liver in vertebrates, fat bodies & follicle
cells in insects)
➢ carried by blood to the ovaries
➢ taken up by microvilli of the PM of the oocytes by
receptor-mediated endocytosis.
*animals may use either method or both (Amphibia)

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 YOLK
• The assembled stored food component = proteins, phospholipids, neutral
fats & glycogen
– Protein yolk (if protein dominates e.g. invertebrates & lower chordates)
– Fatty yolk (if fat dominates)
• Yolk breaks ➔ amino acids, etc. for growing embryo
• Yolk is stored in the form of
– Yolk Platelets (Amphibia & other vertebrates)
– Yolk Granules (invertebrates & lower chordates) &
– Lipochondria (core of accumulated lipid + external protein cover)

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 CONTROL OF VITELLOGENESIS:
Role of Hormones, Nurse cells & Follicular cells
in INSECTS:
Promotes oocyte differentiation
Regulates uptake of yolk proteins at
oocyte surface
Follicle cells→ecdysone→ 20-HE
Juvenile
hormone
Taken in part from Figure 21.12 Gilbert 11th edn.

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 in INSECTS:
2 types of accessory cells play major role in extraovarian transport
of yolk
Follicle cells & Nurse cells (abortive oocytes)
cluster of oocyte + follicle cells + nurse cells = Egg chamber
Follicle cells collect Vitellogenin from blood
& transport it to the egg through the
intercellular gaps & through their microvilli
by pinocytosis
Nurse cells deliver mRNA & proteins into
the oocyte An egg chamber
Figure 9.28 (A) Gilbert 11th edn.

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 In Vertebrates (amphibians): controlled by Estradiol
• Hypothalamus ➔ GnRH
• Pituitary ➔ Gonadotrophins
• Stimulate ovarian follicle to produce estrogen
– helps in oocyte growth
– act on the liver to stimulate vitellogenesis
– by activating the genes for transcription of vitellogenin

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 in INSECTS:

The process of vitellogenesis is

very similar in insects and
vertebrates.
In Vertebrates (e.g. amphibians):
brain stimulates the
hypothalamus (analogous to
corpora allata in insects) to
secrete GnRH which trigger the
pituitary to release
Gonadotrophins. Gonadotrophins,
in turn, stimulate the follicle cells
to produce estrogen, which is the
most potent steroid hormone in
females.
Estrogens helps in oocyte growth &
act on the liver to stimulate
vitellogenesis by activating the
genes for its transcription.

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 in INSECTS:

The process of vitellogenesis is

very similar in insects and
vertebrates.
In Vertebrates (e.g. amphibians):
brain stimulates the
hypothalamus (analogous to
corpora allata in insects) to
secrete GnRH which trigger the

DIY-4
pituitary to release
Gonadotrophins. Gonadotrophins,
in turn, stimulate the follicle cells
to produce estrogen, which is the
most potent steroid hormone in
females.
Estrogens helps in oocyte growth &
act on the liver to stimulate
vitellogenesis by activating the
genes for its transcription.

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 Stages of egg maturation at the time of sperm entry
in different animal species:

Figure 7.3 Gilbert 11th edn.

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 Structure of Ovum - the mature unfertilized egg

• almost all cellular mechanisms

are suspended
• mRNA is masked by proteins
• Minute to Large in size
• Enclosed within the egg
follicles of the ovaries
• Surrounded by protective
membranes except in Sponges,
Coelentrates
Structure of a sea urchin egg
Figure 7.4 Gilbert 6th edn.

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 Corona radiata

Zona pellucida

Perivitelline space

Plasma membrane

Germinal vesicle

Germinal spot/nucleolus

Ooplasm/germplasm

Structure of a mature human ovum

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 • Cytoplasm and nucleus: a remarkable cytoplasmic storehouse
– Nutritive proteins: yolk proteins
– Ribosomes and tRNA: specialized to make structural proteins &
enzymes for the early zygote
– Messenger RNAs: are available to encode proteins for the early early
zygote
– Various Morphogenetic factors: the transcription factors and
paracrine factors are present that help in differentiation of egg
– There are certain Protective chemicals in the ooplasm: eg
antibodies are found in yolk of some species, others have ultraviolet
filters and DNA repair enzymes
– The nucleus of mature ovum is known as Haploid female
pronucleus

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 • Lets talk about the various EGG MEMBRANES /EGG COATS
AND EXTRACELLULAR ENVELOPE
– The protective egg membranes or egg covers around the
plasma membrane of the oocyte
– on the basis of their origin : they are categorized as
• Primary
• Secondary
• Tertiary

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 1. Primary membranes -
• secreted by the egg or follicle cells during oogenesis
• immediately surround the egg
• b/w the egg PM and follicle cells
• types: Vitelline membrane, Zona Pellucida, Zona radiata, Jelly
envelope

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 1.1 Vitelline membrane
insects, molluscs, fish (=called chorion), amphibians, birds
• non cellular, transparent
• Mucoproteins: tensile &
elastic
• Species-specific
recognition of sperm
• fluid filled space b/w the
PM and vitelline
membrane called:
Perivitelline space Sea urchin egg cell surfaces.(A) Scanning electron micrograph of an egg before
(=Vitelline envelope) fertilization. The cell membrane is exposed where the vitelline envelope has been torn.
(B) Transmission electron micrograph of an unfertilized egg, showing microvilli and cell
filled with a fluid called membrane, which are closely covered by the vitelline envelope. A cortical granule lies
perivitelline fluid directly beneath the cell membrane.
Figure th edn.
Course: Developmental Biology Created by:7.4 (b) Gilbert
Dr.Priya Goel, 11
Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 1.2 Zona Pellucida : Mammalian egg only (in non-mammals, it is called Vitelline membrane)
• non cellular, thick
• Glycoproteins (e.g. ZP1, ZP2, ZP3* in
mouse, + ZP4 in humans)
• species-specific recognition of sperm &
egg during fertilization
• induces Acrosome reaction in ZP
bound spermatozoa
• prevents polyspermy Hamster egg
• protects the embryo until implantation Figure 7.5 Gilbert 11 edn. th

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
1.3 Zona Radiata
• fish, some amphibian & reptiles
• Striated appearance
• Present inner to the Vitelline membrane
• Represents the degraded microvilli of the growing oocyte

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 1.4 Jelly Envelope
• marine invertebrates
(Echinoderms: sea urchin)
• Keeps the eggs glued together
• releases fertilizin–attract sperms
of same species
• Resact - Chemotactic molecules
• Helps in attracting & activating Structure of the sea urchin egg at fertilization
Figure 7.2 Gilbert 11th edn.
the sperm

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 2. Secondary membranes:
• secreted by follicle cells around the oocyte, before the egg leaves the
ovary
• outside the primary membrane
• tough, impermeable
• Micropyle: an opening on secondary membrane to facilitate sperm entry

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 2.1 Chorion: chitinous shell (eggs of insects, ascidians and
cyclostomes) outside the vitelline membrane.
- thin in insects which lay eggs in water - acts like a physical gill.
- quite thick in insects which lay eggs in the open air.
- one or more openings called micropyles through which the
sperms enter the egg

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 2.2 Corona radiata (mammals):
Corona radiata

– innermost layer of cumulus cells, Zona pellucida

next to zona pellucida Perivitelline space

Plasma membrane
– Glued to surface of egg through Germinal vesicle

Hyaluronic acid (digested by Germinal spot/nucleolus

hyaluronidase of sperm acrosome) Ooplasm/germplasm

Structure of a mature human ovum

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 3. Tertiary membranes:
• secreted by oviduct/other extraovarian tissues- after ovulation
3.1 Jelly coat: amphibians
• holds the eggs together, swells after fertilization: persists
• provides buoyancy: air bubbles are present
• protection from
– mechanical stress (anchorage to plants)
– predators
– sun UV rays (melanin)
• keep the eggs warm Frog early embryos covered in jelly coat
Course: Developmental Biology Created by: Dr.Priya Goel, Departmenthttps://commons.wikimedia.org/wiki/File:Frog_eggs.jpg
of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 3.2 Albumen : Elasmobranchs
+ horny capsule is secreted by shell glands of oviduct
– corners of the capsule → four long twisted elastic filaments:
attach eggs to sea weeds
– mermaid’s purse/devil’s purse

A Skate Egg Case: a mermaid’s purse

https://commons.wikimedia.org/wiki/File:Mermai
dpurse.jpg

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
3.3 Albumen + Shell membranes + Shell: Reptiles & Birds
• Albumen/Ovalbumin: secreted by glandular wall of oviduct
✓ Water + protein (egg white)
✓ ovum rotates as it passes through the duct
✓ albumen gets wrapped in layers
✓ Chalazae: opaque twisted cords
• Shell membranes: Keratin,
enclose air space at one end
• Shell: CaCO3, laid by shell
gland of oviduct, porous,
collagen Cleavage begins here
Figure 12.2Upadhyaya th edn.
Gilbert 11College
Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal (University of Delhi)
 Functions of egg coats
• Egg membranes protect the eggs & early embryos from
Predators, Dessication, Mechanical stress, Chemical stress,
Ecological hazards (variable temp, pH, radiations, pollution etc.)
• Give buoyancy
• Species-specific sperm attraction & recognition
• Block polyspermy
• Albumen acts as source of nutrition
• Helps in anchorage to sea weeds (mermaid’s purse) & to form egg
mass
Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 Functions of ovum

• carry the set of female chromosomes to restore diploidy

in the zygote and create the right environment to enable
fertilization by the sperm
• acts as a store of cytoplasmic enzymes, mRNAs,
organelles, and metabolic substrates for the growing
embryo until it implants itself into the uterus and the
placenta is formed
• supplies the energy required for cell replication and
division
Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 More about ovum……

• Ovum polarity
• Types of ovum

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 Accumulation of Yolk gives Polarity to the Egg

• Young oocytes: centrally located

germinal vesicle & uniformly
distributed yolk platelets in the
cytoplasm
• Later, internal displacement
introduces polarity:
– germinal vesicle → animal pole
– yolk platelets →vegetal pole
A diagram of an oocyte with its animal and
vegetal hemispheres identified
https://commons.wikimedia.org/wiki/File:Oocyte_Poles.jpg
Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 • POLARITY: The existence of a definite animal- An imaginary polar axis
vegetal axis in the egg , as indicated by the position
of nucleus and yolk
• Animal Pole: region of high activity, polar body gets
pinched off here, nucleus, clear cytoplasm
• Vegetal Pole: yolk-rich cytoplasm (DEUTOPLASM) +
pigment granules
• Polar Axis: the imaginary line passing through
animal & vegetal poles
• Egg polarity also determines the early cleavages in
the zygote

A diagram of an oocyte with its animal and

vegetal hemispheres identified
https://commons.wikimedia.org/wiki/File:Oocyte_Poles.jpg
Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 Types of ovum/egg
based on yolk content
based on yolk distribution
based on the presence/absence of shell
based on the type of development

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 Types of Ovum/Egg
based on Yolk content

ALECITHAL – LECITHAL –
yolk absent or minute Yolk present
(eutherian mammals: humans)
Micro-/ Oligo-lecithal
Small or negligible amount of yolk
Mesolecithal
(Echinoderms,
moderate Amphioxus,
amount ofmarsupials
yolk )
Macro-/Mega-/
(petromyzon, Poly-lecithal
dipnoi, amphibians)
large amount of yolk
cytoplasm-thin cap over the yolk
(insects, reptiles, birds, monotremes)
Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 TYPES OF LECITHAL EGGS
based on yolk distribution
HOMOLECITHAL/ ISOLECITHAL HETEROLECITHAL / ANISOLECITHAL
Uniform distribution of yolk Localized yolk
in ooplasm
(Echinoderms, Amphioxus, Telolecithal
mammals) Yolk in vegetal
Meiolecithal/ half ofTelolecithal
Extreme ooplasm
Yolk occupies (amphibians)
nearly entire ooplasm
Centrolecithal
Disc-like cytoplasm
Yolk is localized for nucleus
in the centre,
(reptiles,thin
Cytoplasm: birds, prototherians)
layer on the surface
+ islands of cytoplasm around nucleus
(insects)
Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
TYPES OF OVUM DEPENDING UPON THE AMOUNT & DISTRIBUTION OF
YOLK

NUCLEUS YOLK ISLANDS

OLIGO/
(placental mammals-
Echinoderms, sea urchin,
humans, monkeys, rodents)
marsupials

MACROLECITHAL
CENTROLECITHAL
annelida, mollusca, reptiles, birds, monotremes
dipnoi, amphibians insects
Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 Types of ovum
on the basis of shell +/-

CLEDOIC EGG NON-CLEDOIC EGG

(insects, reptiles, birds, (All viviparous animals and in
prototherians) oviparous animals
which lays eggs in water)
• Shell: thick, hard • shell: absent
• YOLK makes egg self-sufficient • Not self-sufficient
• CLOSED SYSTEM • Embryo depends on
maternal body for development

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 Types of ovum
on the type of development

DETERMINATE/ INDETERMINATE/
MOSAIC EGG REGULATIVE EGG
Fate of each part of the Fate of each part of the
egg is fixed egg is not fixed
Any blastomere removed Normal development takes place,
experimentally, results in even if any part is removed
the formation of early in development
a defective embryo (echinoderms, chordates)
(annelids, Molluscs, ascidians)
Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 Control of Gametogenesis
Control of spermatogenesis
& oogenesis

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 Control of Spermatogenesis

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
Control of Spermatogenesis

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
Control of Spermatogenesis

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
Control of Spermatogenesis

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 Control of Oogenesis

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
Control of Oogenesis

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
Control of Oogenesis

Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 SUMMARY
• Primordial germ cells (PGCs) –the precursors of gametes- migrate to
gonads in the early embryogenesis.
• Gametogenesis involves 3 phases - Multiplication/Proliferation,
Growth & Maturation - common to spermatogenesis & oogenesis
• Sperm is the motile gamete while ovum is a quiescent, inert cell
with almost all cellular mechanisms are suspended
• Surrounding the egg are a number of protective, functional layers
• Eggs are of many types w.r.t. content and distribution of yolk

Distributed under Creative Commons Attribution-

Non
Course: Developmental Commercial-Share
Biology AlikeGoel,- Department
Created by: Dr.Priya CC-BY-NC-SA.
of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 Time to ponder upon….

➢ Why is gamete formation an essential phase in the cycle of life?

➢ Why is it that spermatogonia behave as stem cells but not
oogonia?
➢ Which great researchers and philosopher(s) is/are credited with
the discovery of mammalian sperm & egg?
➢ What are the essential similarities and differences between the
processes of Spermatogenesis & Oogenesis?
➢ Why is ovum bound to carry that extra burden of ‘dowry’ for the
zygote and not sperm?
Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)
 SUGGESTED READINGS
• Gilbert, SF* and Barresi, MJF (2016) Developmental Biology. XI
Edition. Sinauer Associates, Inc., Publishers, Sunderland,
Massachusetts, USA. ISBN : 9780878939787
• Gilbert SF. (2000) Developmental Biology. IX Edition. Sinauer
Associates, Inc., Publishers, Sunderland, Massachusetts, USA.
ISBN : 9780878939787
• Wolpert, L and Tickle, C (2011) Principles of Developmental
Biology (4th edition). Oxford University Press, ISBN:
9780198792918
• Kalthoff (2008). Analysis of Biological Development, II Edition,
McGraw-Hill Publishers
*all diagrams used herein from this book are with kind permission of Prof. Scott Glibert
Course: Developmental Biology Created by: Dr.Priya Goel, Department of Zoology, Deen Dayal Upadhyaya College (University of Delhi)