CHAPTER 3: GAMETOGENESIS

Gametogenesis
Conversion of germ plasm into highly specialized sex cells (ova & spermatozoa) capable of uniting at fertilization
and producing a new being
Germ Plasm
Gametes + cells that give rise to them
Somatoplasm
Constitutes the cells (somatic), which has no direct part in the production of gametes. It protects
and nourishes the germ plasm
Four Major Phases:
The origin of the germ plasm and their migration to the gonads
The multiplication of the germ cells in the gonads through mitosis
The reduction of the number of chromosomes by one-half by meiosis
The final stages of maturation and differentiation of the gametes into spermatozoa or ova

The Origin of Primordial Germ Cells and their Migration to the Gonads
Characteristics
Large with clear cytoplasm
Have a high alkaline phosphatase activity in mammals and a high glycogen content in birds
Origin
Epiblast
Reptiles, birds and mammals
Take up temporary residence in extraembryonic tissue then return to the embryo proper
Embryonic Mesodermal Cells
Urodele amphibians
Form through the inductive influence of the ventral endodermal yolk
Location
Germinal Crescent
Birds
Located beyond the future head region of the embryo
Posterior Wall of the Yolk Sac
Mammals
Near the region of the allantois

The Origin of Primordial Germ Cells and their Migration to the Gonads
Colonize Gonads
Birds and reptiles
Migrate through the walls of local blood vessels, enter circulation, penetrate the blood vessels of the
gonads, and settle there
Mammalian germ cells migrate around the wall of the posterior gut and then through the dorsal mesentery
Activate migration of the PGCs is guided by the orientation of extracellular matrix molecules, such
as laminin and fibronectin
Chemotactic influence from the gonads guides the final stage of migration
Teratomas
Bizarre tumors containing scrambled mixtures of highly differentiated tissues, which are formed by the
primordial germ cells when they die
PGCs
Increase in number as they migrate to the gonads
Once settled, they enter a proliferative phase in which their numbers increase greatly

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Proliferation of Germ Cells by Mitosis
Oogonia & Spermatogonia
Mitotically active germ cells in females and males, respectively
In human female:
2nd-5th months of pregnancy intense mitotic activity
Number of oogonia then falls sharply because of atresia
7th month most oogonia have entered prophase I as primary oocytes
End of proliferative phase
Non-mammalian vertebrates
Capable of mitosis throughout reproductive life cycle
Fishes release thousands of eggs in one spawning
In males (mammals):
Testes always retain a germinative population of spermatogonia
Periodic waves of mitosis produce subpopulations of spermatocytes

Spermatogenesis
The transition from mitotically active primordial germ cells to mature spermatozoa
Three Principal Phases:
Mitotic Multiplication
Meiosis
Spermiogenesis
Spermatogonia
Mitotically active cells within the seminiferous tubules
Intercellular bridges connecting cytoplasm of daughter cells
Result of incomplete cytokinesis
Facilitates synchronous differentiation and division of sperm-producing cells
During spermatogenesis, the cells are closely associated with sertoli cells because of their functions
Types:
Type-A
Stem cell population
Type-B
Arise form type-A; they finished spermatogenesis
Called preleptotene spermatocytes after the final round of DNA duplication

Sertoli Cells
Form the blood-testis barrier to prevent the bodys immune system from destroying the maturing sperm cells
Blood-testis barrier consists of sertoli processes attached to one another by tight junctions
Create an environment that is important in the differentiation of sperm cells
Facilitate the release of the mature spermatozoa
Degrade the residual spermatozoa

Spermiogenesis
Nuclear Condensation
Nucleus begins to lose fluid
Decrease in size
Chromatin condensation
May serve a protective function to mutation of physical damage
Helps streamline the cell by reducing volume
Acrosome Formation
The cytoplasm streams away form the nucleus becoming the sperm head
Golgi complex at the apical end of the sperm head forms proacrosomal granules
Proacrosomal granules merge to become an acrosome

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 Acrosomes settle close to the nucleus and positions itself like a cap over the largest part of the nucleusThe
acrosome corresponds functionally to a lysosome and thus contains lysosomal enzymes (especially
hyaluronidase)
Development of the Flagellum
Centrioles become a point of anchorage for the developing flagellum
Microtubules from centrioles become continuous with microtubules of flagellum
Mitochondria begin to form a spiral investment around the proximal art of the flagellum
The excess cytoplasm, known as residual bodies is phagocytosed by Surrounding Sertoli Cells in the testes
Sperm Maturation
Newly formed mammalian spermatozoa are minimally functional, unlike invertebrates
Exposure to the environment of the male genital duct system during its transport to the epididymis gives
sperm some degree of motility
Spermiation
The mature spermatozoa are released from the protective sertoli cells into the lumen of the seminiferous
tubule
Removes the remaining unnecessary cytoplasm and organelles
The resulting spermatozoa appears mature but lack motility, rendering them sterile
The non-motile spermatozoa are transported via the testicular fluid secreted by the sertoli cells to the
epididymis where they will acquire some degree of motility
Transport is aided by peristaltic contraction not by the sperms motility
Head of the sperm becomes covered by a glycoprotein
The glycoprotein coat over the acrosome prevents the sperm from fertilizing the egg prior to travelling
through the male and female reproductive tracts
Capacitation of the sperm by the enzymes FPP (fertilization promoting peptide, produced by the male) and
heparin (in the female reproductive tract) remove this coat and allow sperm to bind to the egg

Oogenesis
Varies with the reproductive habits of the animal
In external fertilization: hundreds to hundreds of thousands of eggs per spawning
In internal fertilization: usually 1; seldom > 15 ova
Size of ovum varies
Inside the mothers body: small necessary to store large amounts of materials and yolk in advance
Outside the mothers body: large contains yolky materials needed for the embryos development
Difference in environment dictates difference in egg coverings
Preparation
Maternal genes play a role in guiding and maintaining early embryonic development

Oogenesis in Amphibians
Geared towards the production of large number of eggs
Mitotic phase is not arrested, rather, new batches are produced each year by mitotic division
Requires a 3 year cycle, so the ovary of a mature frog contains 3 batches of eggs everytime

Oogenesis in Amphibians: Development of Amphibian Egg

A developing egg must anticipate the requirements of a developing egg
Supply of yolk
Provides substrates for the synthetic activity of the embryo
The maturing egg synthesizes a large amount of RNA needed for cell division
Three Phases:
Previtellogenic Phase (pre desposition of yolk)
Includes the period up to the early diplotene
Changes prior to diplotene stage:
Increase in the number of mitochondria

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 Increased synthesis of RNA, tRNA, rRNA
Changes during diplotene:
The nucleus (germinal vesicle)
Becomes a site of intense synthetic activity
Increase in diameter up to 7 or 8x
Loops start to form among chromosomes Lampbrush chromosomes
Areas in which DNA are stretched out
RNA synthesis occurs along the loop
Approximately 5% of the genome is exposed and act as template for RNA
synthesis
Formation of large numbers of nucleoli
Associated with specific gene amplification
Adaptive response for meeting the synthetic requirements of
the egg
Formation of ribosomes enough to last the period of cleavage
Vitellogenic Phase (period of yolk deposition
Includes changes that occur in the cytoplasm
Formation of yolk
Most important inclusion
Produced by the liver under the influence of estrogens and carried to the
ovary via the blood
Vitellogenin yolk precursor In the blood
Incorporated in the blood through micropinocytosis
Represented by 2 molecules:
Phosphovitin
Lipovitellin
Yolk nucleus (Balbiani body) migrate to the subcortical region at the vegetal pole
Involved in transforming this region to germ plasm
Cortical granules form
Originate from golgi complex
Dispersed around the cortex of the egg, beneath the plasma membrane
Important in the reaction of the egg and the penetration of sperm cells
Late diplotene or early diakineses
Oocytes are mature
Chromosomes lose their lampbrush configuration and begin to condense near
the center of the nucleus
Nucleoli move from the are adjacent to the inner nuclear membrane to the
center
Prerequisite to the dissolution of the nuclear membrane
With large number of mitochondria
Other cytoplasmic organelles:
Centrioles
Golgi body, for final packaging of carbohydrate and proteins
Special form of ER (annulate lamellae)
Pigment granules
Tentative polarity is formed
Final Maturation (release of oocyte from meiotic block)
Consists of the following stages
Hormonally induced release of the eggs from its first meiotic block
Breakdown of the germinal vesicle
Completion of the first meiotic division
Formation of the first polar body

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 Gonadotropin-induced secretion of progesterone by the follicular cells surrounding the
oocytes
Progesterone acts upon an the surface of the oocyte, causing an electrical depolarization
of the plasma membrane
Results:
Increase in Ca and decrease in cAMP
Production of MPF (Maturation Promoting Factor) which stimulates the transition of
oocyte from G2 to M Phase
RNA synthesis is shut down
Egg is continuously exposed to progesterone causing germinal vesicle to break down
Nucleoplasm and cytoplasm mixes
Egg undergoes meiotic block at metaphase II due to the action of CSF (cytostatic factor)
Sperm penetration releases the mature egg from the CSF induced inhibition
2nd meiotic division is completed
2nd polar body released

Oogenesis in Birds
Yolk
Accumulates within the cytoplasm of the ovum before it is liberated from the ovary
Not synthesized in the ovum but produced in the liver and transported via the blood to the follicular cells
The other components of the egg (shell membrane and shell itself) are non-cellular secretions
Rate of size increase of the ovum accelerates greatly because of yolk accumulation
As more and more yolk accumulate, they crowd toward the surface and project from it
The protuberance containing the ovum is the ovarian follicle
When full allotment of yolk has accumulated in the ovum, ovulation occurred
Ovulation occurs by the rupture of an avascular band, stigma, that surrounds the follicle
Contraction of smooth muscle releases the ovum from the follicle
Like in mammals, completion of MD I is almost in time with ovulation and MD II does not occur without
sperm penetration

Oogenesis in Mammals
Mammals do not replenish their store of oocytes
At birth, human ovaries contain about 1M oocytes arrested at diplotene stage of MD I
Only about 400 of these will reach maturity; the rest will undergo atresia
Mammalian egg contains lipid droplets and glycogen molecules
In humans one egg per month undergoes maturation and ovulation
Oocytes first become associated with follicular cells in the late fetal period
3 Major Phases of Follicle Cells
Primordial Follicle. This will develop into primary follicle
Oocytes at this point are in a period of arrest (1st)
Oocytes and follicular cells develop microvilli connected by gap junctions
Zona pellucida is produced
Growth of both the oocyte and its granulosa cells under the influence of hormones from the pituitary
Granulosa cells actively proliferate
Theca folliculi begins to form
A basement membrane, the membrane granulosa, forms between the granuosa and the overlying
theca
Inner part of the theca becomes vascular
A fluid-filled space, the anrtum, is formed within the granulos cells
At this point, the follicle is referred to as a secondary follicle
Theca cells develop LH receptors on their surface
Thecal cells begin to produce testosterone

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 Granulosa cells by this time have synthesized the enzyme aromatase which converts testosterone
to estrogen within the granulosa cells
Dramatic growth in the population of follicles cells and the final selection of only one follicle that will
undergo ovulation
Begins late in the secretory phase of the menstrual cycle preceding ovulation
The largest of the growing follicle increase greatky in size both by an increase in the liquor folliculi
and by an increase in the number of granulosa cells
Granulosa and theca cells acquire the ability to bind LH
Midcycle LH surge stops
Proliferation of granulosa stops but antral fluid continues to increase
Synthetic activities of angiogenic factors causes granulosa cells of maturing follicles to produce
estradiol
Dominant follicle secretes a factor that blocks the effect of gonadotropins on other follicles and
thus maintains its dominance
A mature graafian is formed
Just before ovulation, the ovum is released from its 1st meiotic block, finishes MD, and releases the
1st polar body
The follicle is now ready to respond to preovulatory LH and FSH surge

Atresia Follicles
The degeneration of the other follcicle cells is called the follicular atresia
The follicle involved is the atretic follicle
Atretic follicles are deficient in receptors for gonadotropins or estradiol
Cells of both the stratum granulosm and the theca interna become involved in the formation of the corpus luteum
Becomes an endocrine organ, secreting both estrogen and progesterone

Accessory Covering of Eggs in Sea Urchins

Adjacent to the plasma membrane is the vitelline envelope
The vitelline envelope of a tough membrane composed of several varieties of glycoproteins
Surrounding the vitelline envelope is the jelly coat
The jelly coat has a high concentration of fucose sulfate polysaccharides
When the eggs are shed, the jelly coat hydrates water and expands

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