GAMETOGENESIS

Gametogenesis lectures will demonstrate many of the important phenomenon we'll

discuss throughout the course - cell-cell interactions, stem cells, hormonal regulation,
differentiation, establishment of polarity. Specializations of gametes reflect their
functions - eggs must provide instructions and nourishment, sperm must deliver the
paternal genetic material. Generally, sperm are among the smallest of cells and eggs
are among the largest. However, a wide variety of cell types exist in both forms of
gametes, especially for eggs.
ells destined to form gametes are set aside very early in development. !hese are
called the PRIMORDIAL GERM CELLS "#G$. !he primordial germ cells are not
initially located in the gonads, but migrate there from other areas of the embryo. !he
#Gs of both sexes appear identical.
#Gs are diploid and must undergo meiosis. %n sperm differentiation meiosis occurs
early and the cells differentiate with haploid nuclei& in oocyte differentiation meiosis
occurs very late, with meiosis %% occurring in some species after fertilization.
!here is a fundamental division of function between germ cells and somatic cells -
only the germ cells will contribute to a new generation& all somatic cells are fated to
death.
SPERM EGG
small in size large in size
continuous production "in most mammals$
produced early in development "in most
mammals$
relatively little interspecific variation relatively large interspecific variation
differentiation occurs after meiosis differentiation occurs before meiosis
motile non-motile
condensed nucleus normal nucleus
SPERMATOGENESIS
Sperm are highly specialized for the delivery of the paternal genome to the egg. !he
tail is a flagellum containing the AXONEME, a structure composed primarily of
microtubules in a '()*' array and tangential and radial arms composed largely of
dynein +. ,n extremely large number of mitochondria are in the middle piece of the
tail, the highest concentration of any cell.
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Spermatogenesis illustrates the principle of critical cell-cell interactions during
development. Sertoli cells critical for spermatogonia survival and differentiation-
spermatogonia cannot be grown in isolation in tissue culture& if co-cultured with
Sertoli cells, they proliferate and produce sperm.
Sperm differentiate within seminiferous tubules of the testes. .ore mature sperm are
found on inside of tubule and less mature near periphery "basal lamina$.
Spermatogonia are STEM CELLS. , stem cell is a cell that divides to replenish itself
and provide a cell that is committed for differentiation. Stem cells are critical in many
developmental processes, including the generation of cells of the blood and immune
system, epidermis, and intestinal epithelium.
Sperm cell differentiation primarily occurs after meiosis. Strange - wouldn't this be a
problem since the haploid genome will express many deleterious recessive mutations
and because half the spermatozoa would lac/ the essential 0 chromosome1 Sperm
circumvent this problem by remaining connected through cytoplasmic bridges during
their differentiation. !hey form a syncytium, a common cytoplasm holding multiple
nuclei. !his allows the exchange of essential gene products. !herefore, sperm really
differentiate using the diploid, not the haploid genome.
2vidence derives from the examination of mutants, specifically dis3unction-defective
mutants of Drosophila that fail to divide chromosomes e4ually between cells at
meiosis. %n these strains, even when sperm contain no chromosomes, differentiation is
normal.
56, in sperm head is highly condensed. Histones are replaced by small highly basic
proteins /nown as protamines. , conse4uence of condensation is that transcription is
shut down in the terminal stages of spermiogenesis. Some new proteins activities
must be appear. How can this happen1 %t suggests that the control of differentiation
must be exerted at the post transriptional le!el"
!he final steps of differentiation occur after release of sperm. Sperm become motile
only after transit through the epididymis. !hey become capable of fertilization only
after residing in the vaginal tract where factors interact with the sperm membrane.
!his process is /nown as CAPACITATION.
#ORMONAL REG$LATION % Spermatogenesis is dependent on androgens "male-
specific steroid hormones$, especially testosterone. ,ndrogens are produced by the
7eydig cells upon stimulation by luteinizing hormone. !hese direct the Sertoli cells to
influence the sperm differentiation.
OOGENESIS
2ggs are remar/able - within days or wee/s of fertilization they give rise to a new
individual. 2ggs are the only cell capable of this. 2ggs have multiple functions that
include- nourishment of the embryo and provision of preformed macromolecular
materials provide pattern information to the embryo.
*
2ggs vary widely in size and complexity among species. !his variety can be
understood at least partially by loo/ing at the comparative strategies of development -
amphibians and fish lay eggs in water and produce free-living larval or 3uvenile forms
relatively early, therefore, eggs typically are of moderate size& birds and reptiles
produce much larger 3uvenile forms on land, therefore eggs are very large to provide
for more nutrient reserves, and have more complex coverings to prevent desiccation
while allowing gas exchange. .ammalian eggs are tiny by comparison - again
understandable because of maternal provision of nutrients.
8ocyte specializations constrain patterns of cleavage. leavage refers to the initial
cells divisions following fertilization. leavage is slower through yol/-rich regions of
eggs. !herefore, cleavage patterns are very different in isolecithal "sparse yol/&
mammals$, mesolecithal "intermediate amounts of yol/& some fish, amphibians, and
echinoderms$, telolecithal "heavy, polarized distribution of yol/& birds and reptiles$,
and centrolecithal "heavy yol/ concentration in center& insects$ eggs.
!he first embryonic axis is often established by the distribution of components in the
egg. !his first may be the anterior-posterior axis "in amphibians, for example$ or the
dorsoventral "birds, for example$ depending on the species.
,ll these points ma/e it clear that oogenesis sets fundamental patterns and constraints
on the developmental program.
2gg development is long and complex. 8ne example is provided by human oogenesis.
8nly a small fraction of the oogonia ever develop to oocytes - the ma3ority die before
birth. 9urthermore, there is a long period of dormancy before final differentiation. 9or
some oocytes, this is as long as :; years.
!he timing of meiosis is very different in oogenesis and spermatogenesis. .eiosis is
completed after most differentiation occurs in oocytes, and before the ma3ority of
differentiation in sperm.
yto/inesis is very different too - eggs conserve cytoplasm by forming polar bodies,
while sperm divide cytoplasm evenly. !his can be understood by the need to conserve
precious resources for early nourishment of the embryo.
.any eggs contain massive reserves of nutrients and macromolecules such as
m<6,s, t<6,s, and ribosomes. %n some organisms, notably amphibians, the large
excess of particular components, especially r<6,s and t<6,s, is due to
amplification of the cognate genes.
Special mechanisms must be employed to achieve rapid oocyte growth. 9or example,
typical somatic cells are +;-*; um in diameter and double in =*> h, yet eggs of
insects that live only days may be +;;; um in diameter, a +;
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increase in mass over
the somatic cell. How can eggs become so large1 !hrough help from ACCESSOR&
CELLS. 2ggs develop in tight association with other cells /nown as accessory cells.
6ote that this is another example of a cell-cell interaction in development.
@
!here are two types of oocyte accessory cells - nurse cells and follicle cells. 6urse
cells are found primarily in insects& follicle cells occur in all organisms, including
insects. 6urse cells are derived from the germ line and remain connected as a
syncytium "multiple nuclei within a common cytoplasm$ with the oocyte. 9ollicle
cells are somatic and don't usually have direct cytoplasmic connections. ,ccessory
cells mediate hormonal interactions, provide nutrients, and in the case of nurse cells,
provide macromolecules to the oocyte.
6urse cells arise from mitotic division of a STEM CELL" <ing canals connect cells -
these allow passage of macromolecules. 8nly one cell of the syncytium develops into
an oocyte, the others form nurse cells. 6urse cells undergo further mitoses without
cyto/inesis to become highly POL&PLOID" !his aids them in the rapid synthesis of
components re4uired for oogenesis. #reformed macromolecules pass from nurse cells
to the oocyte.
!he movement of material from the nurse cell to the oocyte is important not only for
building nutritional reserves, but also for laying down instructions for the embryo's
body plan. .any substances that define embryonic pattern are transported from the
nurse cells and localized within the oocyte during this period.
'OLLICLE CELLS do not feed large macromolecules directly to the oocyte, but
mediate the transport of many material from the blood or hemolymph to the oocyte
and are /ey sources of hormones re4uired for oocyte maturation.
2gg cell structure is highly complex. !he nucleus "also /nown as the GERMINAL
(ESICLE) generally is large and eccentrically placed. %n all but isolecithal eggs,
there is a polarized distribution of yol/, ribosomes, mitochondria, and often pigment
granules. !he relatively yol/-free region of the oocyte is /nown and the ANIMAL
#EMISP#ERE and the yol/-rich region is /nown as the (EGETAL
#EMISP#ERE. !he establishment of the animal-vegetal axis "the imaginary line
connecting the two poles of each hemisphere$ defines the anterior-posterior axis of the
amphibian embryo, the first specification of the body plan. !he oocyte cytoplasm is
divided into a CORTICAL AND S$*CORTICAL REGIONS.
!he cortex is gel-li/e, rich in cytos/eletal elements, and contains membrane enclosed
organelles "CORTICAL GRAN$LES) that are important in creating the fertilization
membrane.
2gg coats play a role in protection and species specific recognition. #rimary egg coats
are elaborated by the oocyte, and include the +ONA PELL$CIDA of mammals and
the (ITELLINE LA&ER of amphibians. Secondary coats are synthesized by the
follicle cells and include the chorion of insects. 9inally, in some eggs, a tertiary coat is
added during passage through the oviduct. 8ne example is the albumin "egg white$
and shell of avian eggs.
&OL, is made in the fat bodies of insects and liver of vertebrates and shunted
between the circulatory system and the egg by the follicle cells. !his obviates a need
for the oocyte to produce large 4uantities of this primary nutrient.
, discussion of the hormonal control of ovulation is beyond the scope of this course.
%t is, however, important to realize that oogenesis is under tight and complex
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hormonal control in all species. 9ollicle cells play an important role in mediating the
interaction between the oocyte and the endocrine system.

CONCL$SIONS
+$ .any important principles of development, including cell-cell interaction, the use
of stem cells, hormonal influences, and differentiation, are demonstrated during
gametogenesis.
*$ 5ifferences in gamete specialization can be understood on the basis of different
re4uirements for development.
@$ %n humans, sperm are produced continuously and eggs are produced only early in
development.
>$ Sperm cells are highly specialized for the delivery of the paternal genome.
:$ Sperm maturation depends on interaction with Sertoli cells and hormones produced
by 7eydig cells.
?$ 2ggs are specialized for their role in nourishment and for the provision of
macromolecular materials and developmental determinants.
A$ ,ccessory cells are essential for oogenesis because they provide a source of
nutrients, hormones, and developmental determinates. 9ollicle and nurse cells are the
two forms of accessory cells.
B$ 2ggs often employ special mechanisms for rapid growth.
($ , complex interplay of hormones triggers and directs oogenesis.
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