Gametogenisis (Course)

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ANATOMY OF DEVELOPMENT

Follows the development of body from procreation to death (ontogenesis) but also
the changes in the historical development of the species (phylogenesis).

Ontogenesis is defined as that seeks to find specialty anatomic and functional


morphological development and transformation of body.
Contains 2 parts:
1. fecundation of birth (prenatal or intrauterine)
2. from birth to death (postnatal)

Human Embryology designed micro and macroscopic morphogenesis of organs and


tissues.
Consistent with the chronological development, is 3 parts of study:
reproduction, deals with the study of gametes: gametogenesis and
fecundation,
embryogenesis, embryonic period, is the study of fecundation by the end
of the week 8th,
organogenesis, fetal period, from the end. S8 - birth

Reproduction and embryo are organized in general embryology and the


organogenesis is the special embryology.
SEXUAL REPRODUCTION

Human development begins by fertilization when the male and


female gametes unite to form a single, unicellular organism, zygote.
Human beeing, as the most living systems of nature, presents
sexually dimorphic adult organisms, whose most important biological
function is sexual reproduction; they are dedicated in producing the next
generation of individuals. Sexual reproduction involves exchanges of the
genetic material, rezulting individuals similar to their parents and a diversity
of individuals of the same generation. A real diversity may be ensured by
avoiding the consanguinity by incest marriages.
Extensive imbreeding increases the frequency of excessive
mutations and congenital birth defects.
Main Features of Gametogenesis

Gametogenesis is the process of cytodifferenciation that converts the germinal line


cells into mature male and female gametes. It assures preservation of chromosomes
number. The earliest stage of the development of a new individual involves mergying of two
gametes, oocyte and spermatozoon.
This represents the essence of the sexual reproduction and is called fecundation
(fertilization, amphimixis). The somatic cells of an adult organism contain a diploid set of
chromosomes: 44+XY for male and 44+XX for female.
They contain the deoxyribonucleic acid (DNA) that encodes all informations
required for the development and function of a organism. In fact, there are 22 pairs of somatic
chromosomes and thus two complete copies of the genetic material of the organism. If the
gametes would be also diploid, the number of chromosomes would double with each
generation. Therefore, it becomes necessary a mechanism for dividing the genetic material.
That happens during the reduction division of meiosis.
Meiosis characterizes only the germinal line in which germinal cells divide and
increase in number, with a simultaneously reduction by half of their number of chromosomes.
The result of meiosis is a haploid gamete from a diploid cell. Each haploid gamete contains
half of the number of chromosomes and each chromosome contains only n DNA. During
fertilization two haploid gametes unite to form a single zygote, thus conserving the 2n DNA
and chromosome number from generation to generation.
An important aspect of meiosis is the exchange of genetic material between
homologous chromosomes, which eventually results in variability in the descendents of a
single pair of parents ensuring genetic diversity.
Furthermore, gametogenesis involves production of specialized sexually
dimorphic gametes that are trully different.
By fertilization a male and female haploid gamete create a single zygote with a
comlete set of chromosomes, half of them paternal and the other half maternal.
This single diploid structure begins then a spectacular developmental history
which culminates in formation of a new adult organism reproductively mature.
This process is called ontogenesis.
Behaviour of Sex Chromosomes during Gametogenesis

Female and male gametes show morphologic differences. In higher


mamals, including humans, female produces only one kind of gametes. Each of
them contains a haploid set of somatic chromosomes (autosomes) and one kind of
sex chromosomes (gonosomes), the X-chromosome. The male produces
spermatozoa of two different kinds. All of them have a haploid set of autosomes, but 50%
have an X-chromosome and 50% have a Y-chromosome. Two notions often confuse
must be emphasized.
Ploidy refers to the number of copies of each chromosome present in a cell
nucleus whereas the n number refers to the number of copies of each unique DNA
strand.
Replication is the process of duplication of both ploidy and n copies at
different stages of the cell cycle (whether mitotic or meiotic).
The genetic sex of the organism is determined at fertilization. The semen's
high sperm density forms a statistically significant population, practically infinite that
ensures an equal chance for both XX or XY combinations. When an oocyte is
fertilized by an X-sperm an XX-zygote (female) will form. When an ovum is
'fertilized by a Y-sperm results a XY-zygote (male). Thus, a male and female zygotes
are alike in respect to their content of autosomes, but differ in respect to their content of
gonosomes. The behavioural traits associated with maleness or femaleness are
determined later during development.
MITOSIS

Mitosis is the process whereby one cell divides, giving rise to two daughter cells
that are genetically identical to the parent cell. Each daughter cell receives the complete
complement of 46 chromosomes. Before a cell enters mitosis, each chromosome
replicates its deoxyribonucleic acid (DNA). During this replication phase the chromosomes
are extremely long, they are spread diffusely through the nucleus.
Mitosis division through the five stages: 1.prophase, 2.prometaphase, 3.
metaphase, 4. anaphase and 5. telophase.
With the onset of mitosis the chromosomes begin to coil, contract, and
condense; these events mark the beginning of prophase. Each chromosome now consists
of two parallel subunits, chromatids, that are joined at a narrow region common to both
called the centromere. Throughout prophase the chromosomes continue to condense,
shorten, and thicken (Fig. A), but only at prometaphase do the chromatids become
distinguishable (Fig. B). During metaphase the chromosomes line up in the equatorial
plane, and their doubled structure is clearly visible (Fig. C ).
Each is attached by microtubules extending from the centromere to the
centriole, forming the mitotic spindle. Soon the centromere of each chromosome
divides, marking the beginning of anaphase, followed by migration of chromatids to
opposite poles of the spindle. Finally, during telophase, chromosomes uncoil and
lengthen, the nuclear envelope reforms, and the cytoplasm divides (Fig. D and E ).
Each daughter cell receives half of all doubled chromosome material and
thus maintains the same number of chromosomes as the mother cell.
Meiosis is the cell division that takes place in the germ cells to generate male
and female gametes, sperm and egg cells, respectively. Meiosis requires two cell
divisions, meiosis I and meiosis II, to reduce the number of chromosomes to
the haploid number of 23.
First meiotic division has an extended prophase period with 5 distinct phases:
leptotene, zygotene, pachytene, diplotene and diakinesis.
In this long prophase the DNA duplicate resulting a 4n amount of DNA. After DNA
duplication in each chromosome the cromatids duplicate resulting bivalent chromosomes.
Then, the same pair of chromosomes (one from father and one from mother)
match up centromere to centromere forming chiasma and exchange large segments of
DNA. This process of genetic recombination is called crossing-over and ensures
genetic variability.
The first meiotic metaphase, anaphase and telophase are characterized by
separation of bivalent chromosomes resulting two haploid cells with 2n DNA chromosoms
(each with two cromatids).

The second meiotic division involves the bivalent chromosomes, whose


centromeres divide and one half of each chromosome passes in each daughter-cell. Thus
results a n DNA amount for each germcell.
First and second meiotic divisions. A. Homologous chromosomes approach each other. B. Homologous
chromosomes pair, and each member of the pair consists of two chromatids. C. Intimately paired
homologous chromosomes interchange chromatid fragments (crossover). Note the chiasma. D. Double-
structured chromosomes pull apart. E. Anaphase of the first meiotic division. F and G. During the second
meiotic division, the double-structured chromosomes split at the centromere. At completion of division,
chromosomes in each of the four daughter cells are different from each other.
The timing of meiosis is different in male and female.
In male, spermatogonia remains in a dormant (sleeping) stage from the 6th week
of the embryonic life until puberty when meiosis appears as the start point for each team
of maturation.
In female, oogonia become oocytes during the 5th month of the fetal
development, enter the first prophase of meiosis and stop in the first meiotic arrest. At
puberty meiosis is taken again (1st day of menstrual cycle) from the first meiotic arrest and
stops in the second meiotic arrest just after ovulation. This process occurs monthly for
every maturation team.
In conclusion, after the two divisions of meiosis, haploid n DNA germ cells
result.
SPERMATOGENESIS .
Male gametogenesis consists of three phases:
Spermatocytosis, when spermatogonia proliferate by mitotic divisions to replace
them- selves and to produce primary spermatocytes. Male meiosis, when, after two
succesive maturation divisions are produced secondary spermatocytes and spermatids, all
haploid n DNA cells. Spermiogenesis, when spermatids become mature sperm cells and
develop movement anexes.

Spermatocytosis
It is a continous process that follows three stages.
Prepubertal stage begins in the 3rd week of the embryonary life when the pri-
mordial sexual stem cells develop from the mesoderm of the yolk sac and migrate to the
posterior body wall by the way of the dorsal hindgut mesentery. At the level of the
mesonephrotic intermediate mesoderme they induce male gonade formation. During
embryonic life these cells divide mitotically and produce spermatogonia. Divisions stop in the
6th week and spermatogonia enter the dormant stage until puberty. Adult stage begins at
puberty and persists throughout the life decreasing after 60 years.
Cellular changes during spermatocytosis. Just prior the puberty spermatogonia
begins to differentiate into primary spermatocytes. Spermatogonia derived from the stem
cell are described as dark A type. They divide mitotically: some of them produce two new
dark A type spermatogonia replenishing thus the complement of germ-cells; others produce
two light A type spermatogonia. Mitotic division of light A type spermatogonia give rise to two
B type spermatogonia. Each B type spermatogonia divides mitotically again resulting two
resting primary spermatocytes.
Male meiosis
Primary spermatocyte replicates its DNA, undergoes the first maturation division
and formes two secondary spermatocytes that contain a haploid number of bivalent (2n
DNA) chromosomes. After a brief interphase each secondary spermatocyte undergoes the
second maturation division without DNA replication. The two resultant cells are spermatides,
haploid cells with monovalent (n DNA) chromosomes.

Spermiogenesis
During this phase spermatides undergo complexe, successive series of changes
to become spermatozoa: reduction of citoplasma; fusion of Golgi complex vesicles to form
a single acrosomal vesicle; fixation of acrosomal vesicle on the nuclear membrane and
edification of future sperm at proximal area.

Coincident with these changes,


cytoplasma migrates to the opposite part of the
cell, pushes the external wall of the acrosomal
vesicle in contact with the cellular membrane and
the spermatide elongates. The two centrioles are
situated near the posterior aspect of the nucleus.
The proximal one remains unmodified; the other
changes into distal or basal body of the sperm
head.
It is the origin of the axial filament and its development contibutes to the sperm
elongation. Around the axial filament mithocondria condense and form a helical sheath.
Development is completed by the increasing of the tail, and the detachement of some
cytoplasma as a residual body. The evolution of the spermatogenetic line cell elements is
accompanied by their gradual translocation from the basement membrane to the lumen of
seminiferous tubes between two adjacent Sertoli cells. Thus the lumen contains only
morphological mature sperm cells. Spermatogenesis Is a continuous process from
puberty to death. It begins in different local areas of the germinal epithelium but it is not
synchronized throughout the seminiferous tubules. All spermatogonia entering the
spermatogenesis in the same time form the team of maturation.
spermatogonium
It begins in different local areas of the germinal
epithelium but it is not synchronized throughout the
seminiferous tubules. All spermatogonia entering the
spermatogenesis in the same time form the team of maturation.
The timing of each human spermatogenesis cycle
is about 64 days as follows: spermatogonial mitosis 16 days;
first meiotic division 8 days; second meiotic division 16
days; spermiogenesis about 24 days.
The origin of the spermatogonia involved in the team of
maturation could be clonal, continual waves of
spermatogenesis occuring allong the seminiferous epithelium.

spermatozoa
Morphology of the sperm

Sperm, the male gamete, is a motile cell free-swiming in the Acrosomal cap

seminal liquid. It consists of a head, neck, body (middle piece) and tail
that ensures its mobility.
The head consists of an acrosomal cap that contains
acrosomase, an enzyme that facilitates the penetration of corona radiata.
A condensed nucleus occupies the most part of the head under the
acrosomal cap. Both are enveloped in a continous plasma membrane,
without intervening cytoplasma.
The neck appears between the head and the middle piece as a
slight constriction. It contains the proximal and distal centrioles.
The body has an axial filament surrounded by a mithocondrial
sheath mithocondria beeing arranged in a helical manner. They ensure
the energy requirements of the sperm.
The tail is the greatest part of the spermatozoon (about 40
microns), a whipelike projection that ensures the sperm movements.
Seminal plasma.The fluid component of the semen contains a
great number of substances, including mucoproteins, proteolytic Endpiece

enzymes, prostaglandines (with important pharmacodynamic actions on


the uterus), acids like citric, uric, lactic, piruvic and sugars as sorbitol,
inozitol and fructose. The later is an essential substance for the anaerobic
glycolysis that ensures sperms survival in the female genital tract.
OOGENESIS

Maturation of Oocytes Begins Before Birth

Once primordial germ cells have arrived in the gonad of a


genetic female, they differentiate into oogonia. These cells undergo a
numberof mitotic divisions and, by the end of the third month, are
arranged in clusters surrounded by a layer of flat epithelial cells.
Whereas all of the oogonia in one cluster are probably derived
from a single cell, the flat epithelial cells, known as follicular cells,
originate from surface epithelium covering the ovary.
The majority of oogonia continue to divide by mitosis, but some
of them arrest their cell division in prophase of meiosis I and form
primary oocytes.
During the next few months, oogonia increase rapidly in number,
and by the fifth month of prenatal development, the total number of germ
cells in the ovary reaches its maximum, estimated at 7 million.
At this time, cell death begins, and many oogonia as well as
primary oocytes become atretic. By the seventh month, the majority of
oogonia have degenerated except for a few near the surface.
All surviving primary oocytes have entered prophase of meiosis I,
and most of them are individually surrounded by a layer of flat epithelial
cells
A primary oocyte, together with its surrounding flat epithelial
cells, is known as a primordial follicle.
Maturation of Oocytes Continues at Puberty

Near the time of birth, all primary oocytes have started prophase of
meiosis I, but instead of proceeding into metaphase, they enter the
diplotene stage, a resting stage during prophase that is characterized by
a lacy network of chromatin.
Primary oocytes remain in prophase and do not finish their
first meiotic division before puberty is reached, apparently because of
oocyte maturation inhibitor (OMI), a substance secreted by follicular
cells. The total number of primary oocytes at birth is estimated to vary from
700,000 to 2 million.
During childhood most oocytes become atretic; only approximately
400,000 are present by the beginning of puberty, and fewer than 500 will
be ovulated. Some oocytes that reach maturity late in life have been
dormant in the diplotene stage of the first meiotic division for 40 years or
more before ovulation.
Whether the diplotene stage is the most suitable phase to protect
the oocyte against environmental influences is unknown. The fact that the
risk of having children with chromosomal abnormalities increases with
maternal age indicates that primary oocytes are vulnerable to damage as
they age.
At puberty, a pool of growing follicles is established and
continuously maintained from the supply of primordial follicles.
Each month, 15 to 20 follicles selected from this pool begin to mature,
passing through three stages:
1) primary or preantral;
2) secondary or antral (also called vesicular or Graafian); and
3) preovulatory.
The antral stage is the longest, whereas the preovulatory stage
encompasses approximately 37 hours before ovulation.
As the primary oocyte begins to grow, surrounding follicular cells
change from flat to cuboidal and proliferate to produce a stratified epithelium
of granulosa cells, and the unit is called a primary follicle.
Granulosa cells rest on a basement membrane separating
them from surrounding stromal cells that form the theca folliculi.
Also, granulosa cells and the oocyte secrete a layer of
glycoproteins on the surface of the oocyte, forming the zona pellucida
As follicles continue to grow, cells of the theca folliculi organize into an
inner layer of secretory cells, the theca interna, and an outer fibrous
capsule, the theca externa.
Also, small, finger-like processes of the follicular cells extend
across the zona pellucida and interdigitate with microvilli of the plasma
membrane of the oocyte. These processes are important for transport
of materials from follicular cells to the oocyte.

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