Spermatogenesis

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Seminiferous tubule with maturing sperm. H&E stain.

Spermatogenesis is the process by which male spermatogonia develop into mature spermatozoa,
also known as a sperm cell. Spermatozoa are the mature male gametes in many sexually
reproducing organisms. Thus, spermatogenesis is the male version of gametogenesis. In
mammals it occurs in the male testes and epididymis in a stepwise fashion, and for humans takes
approximately 64 days.[1] Spermatogenesis is highly dependent upon optimal conditions for the
process to occur correctly, and is essential for sexual reproduction. It starts at puberty and
usually continues uninterrupted until death, although a slight decrease can be discerned in the
quantity of produced sperm with increase in age. The entire process can be broken up into
several distinct stages, each corresponding to a particular type of cell:.

Cell type ploidy/chromosomes chromatids Process

spermatogonium (types Ad, Ap spermatocytogenesis
diploid/46 2N:2C
and B) (mitosis)
spermatidogenesis (meiosis
primary spermatocyte diploid/46 2N:4C
1)
spermatidogenesis (meiosis
secondary spermatocyte haploid/23 1N:2C
2)
spermatid haploid/23 1N:1C spermiogenesis
sperm haploid/23 1N:1C spermiation
A mature human Spermatozoon
Contents
[hide]

 1 Purpose
 2 Location
 3 Stages
o 3.1 Spermatocytogenesis
o 3.2 Spermatidogenesis
o 3.3 Spermiogenesis
 4 Role of Sertoli cells
 5 Influencing factors
 6 Hormonal control
 7 See also
 8 References
 9 External links

[edit] Purpose
Spermatogenesis produces mature male gametes, commonly called sperm but specifically known
as spermatozoa, which are able to fertilize the counterpart female gamete, the oocyte, during
conception to produce a single-celled individual known as a zygote. This is the cornerstone of
sexual reproduction and involves the two gametes both contributing half the normal set of
chromosomes (haploid) to result in a chromosomally normal (diploid) zygote.

To preserve the number of chromosomes in the offspring – which differs between species – each
gamete must have half the usual number of chromosomes present in other body cells. Otherwise,
the offspring will have twice the normal number of chromosomes, and serious abnormalities may
result. In humans, chromosomal abnormalities arising from incorrect spermatogenesis can result
in Down Syndrome, Klinefelter's Syndrome, and spontaneous abortion.

[edit] Location
Spermatogenesis takes place within several structures of the male reproductive system. The
initial stages occur within the testes and progress to the epididymis where the developing
gametes mature and are stored until ejaculation. The seminiferous tubules of the testes are the
starting point for the process, where stem cells adjacent to the inner tubule wall divide in a
centripetal direction—beginning at the walls and proceeding into the innermost part, or lumen—
to produce immature sperm. Maturation occurs in the epididymis and involves the acquisition of
a tail and hence motility.

[edit] Stages
[edit] Spermatocytogenesis

Schematic diagram of Spermatocytogenesis

Main article: Spermatocytogenesis

Spermatocytogenesis is the male form of gametocytogenesis and results in the formation of

spermatocytes possessing half the normal complement of genetic material. In
spermatocytogenesis, a diploid spermatogonium which resides in the basal compartment of
seminiferous tubules, divides mitotically to produce two diploid intermediate cells called primary
spermatocytes. Each primary spermatocyte then moves into the adluminal compartment of the
seminiferous tubules and duplicates its DNA and subsequently undergoes meiosis I to produce
two haploid secondary spermatocytes. This division implicates sources of genetic variation, such
as random inclusion of either parental chromosomes, and chromosomal crossover, to increase the
genetic variability of the gamete.

Each cell division from a spermatogonium to a spermatid is incomplete; the cells remain
connected to one another by bridges of cytoplasm to allow synchronous development. It should
also be noted that not all spermatogonia divide to produce spermatocytes, otherwise the supply
would run out. Instead, certain types of spermatogonia divide to produce copies of themselves,
thereby ensuring a constant supply of gametogonia to fuel spermatogenesis.

[edit] Spermatidogenesis

Main article: Spermatidogenesis

Spermatidogenesis is the creation of spermatids from secondary spermatocytes. Secondary

spermatocytes produced earlier rapidly enter meiosis II and divide to produce haploid
spermatids. The brevity of this stage means that secondary spermatocytes are rarely seen in
histological preparations.

[edit] Spermiogenesis

Main article: Spermiogenesis

During spermiogenesis, the spermatids begin to grow a tail, and develop a thickened mid-piece,
where the mitochondria gather and form an axoneme. Spermatid DNA also undergoes
packaging, becoming highly condensed. The DNA is packaged firstly with specific nuclear basic
proteins, which are subsequently replaced with protamines during spermatid elongation. The
resultant tightly packed chromatin is transcriptionally inactive. The Golgi apparatus surrounds
the now condensed nucleus, becoming the acrosome. One of the centrioles of the cell elongates
to become the tail of the sperm.

Maturation then takes place under the influence of testosterone, which removes the remaining
unnecessary cytoplasm and organelles. The excess cytoplasm, known as residual bodies, is
phagocytosed by surrounding Sertoli cells in the testes. The resulting spermatozoa are now
mature but lack motility, rendering them sterile. The mature spermatozoa are released from the
protective Sertoli cells into the lumen of the seminiferous tubule in a process called spermiation.

The non-motile spermatozoa are transported to the epididymis in testicular fluid secreted by the
Sertoli cells with the aid of peristaltic contraction. While in the epididymis the spermatozoa gain
motility and become capable of fertilization. However, transport of the mature spermatozoa
through the remainder of the male reproductive system is achieved via muscle contraction rather
than the spermatozoon's recently acquired motility.

[edit] Role of Sertoli cells

Labelled diagram of the organisation of Sertoli cells (red) and spermatocytes (blue) in the testis.
Spermatids which have not yet undergone spermination are attached to the lumenal apex of the
cell
Main article: Sertoli cell

At all stages of differentiation, the spermatogenic cells are in close contact with Sertoli cells
which are thought to provide structural and metabolic support to the developing sperm cells. A
single Sertoli cell extends from the basement membrane to the lumen of the seminiferous tubule,
although the cytoplasmic processes are difficult to distinguish at the light microscopic level.

Sertoli cells serve a number of functions during spermatogenesis, they support the developing
gametes in the following ways:

 Maintain the environment necessary for development and maturation via the blood-testis
barrier
 Secrete substances initiating meiosis
 Secrete supporting testicular fluid
 Secrete androgen-binding protein, which concentrates testosterone in close proximity to
the developing gametes
o Testosterone is needed in very high quantities for maintenance of the reproductive
tract, and ABP allows a much higher level of fertility
 Secrete hormones affecting pituitary gland control of spermatogenesis, particularly the
polypeptide hormone, inhibin
 Phagocytose residual cytoplasm left over from spermiogenesis
 They release Antimullerian hormone which prevents formation of the Mullerian Duct /
Oviduct.
 Protect spermatids from the immune system of the male.

[edit] Influencing factors

The process of spermatogenesis is highly sensitive to fluctuations in the environment,
particularly hormones and temperature. Testosterone is required in large local concentrations to
maintain the process, which is achieved via the binding of testosterone by androgen binding
protein present in the seminiferous tubules. Testosterone is produced by interstitial cells, also
known as Leydig cells, which reside adjacent to the seminiferous tubules.

Seminiferous epithelium is sensitive to elevated temperature in humans and some other species,
and will be adversely affected by temperatures as high as normal body temperature.
Consequently, the testes are located outside the body in a sack of skin called the scrotum. The
optimal temperature is maintained at 2 °C (man) - 8 °C (mouse) below body temperature. This is
achieved by regulation of blood flow[2] and positioning towards and away from the heat of the
body by the cremasteric muscle and the dartos smooth muscle in the scrotum.

Dietary deficiencies (such as vitamins B, E and A), anabolic steroids, metals (cadmium and
lead), x-ray exposure, dioxin, alcohol, and infectious diseases will also adversely affect the rate
of spermatogenesis.

[edit] Hormonal control

Hormonal control of spermatogenesis varies among species. In humans the mechanism are not
completely understood, however it is known that initiation of spermatogenesis occurs at puberty
due to the interaction of the hypothalamus, pituitary gland and Leydig cells. If the pituitary gland
is removed, spermatogenesis can still be initiated by follicle stimulating hormone and
testosterone.

Follicle stimulating hormone stimulates both the production of androgen binding protein by
Sertoli cells, and the formation of the blood-testis barrier. Androgen binding protein is essential
to concentrating testosterone in levels high enough to initiate and maintain spermatogenesis,
which can be 20-50 times higher than the concentration found in blood. Follicle stimulating
hormone may initiate the sequestering of testosterone in the testes, but once developed only
testosterone is required to maintain spermatogenesis. However, increasing the levels of follicle
stimulating hormone will increase the production of spermatozoa by preventing the apoptosis of
type A spermatogonia. The hormone inhibin acts to decrease the levels of follicle stimulating
hormone. Studies from rodent models suggest that gonadotropin hormones (both LH and FSH)
support the process of spermatogenesis by suppressing the proapoptotic signals and therefore
promote spermatogenic cell survival. [3]

The Sertoli cells themselves mediate parts of spermatogenesis though hormone production. They
are capable of producing the hormones estradiol and inhibin. The Leydig cells are also capable of
producing estradiol in addition to their main product testosterone.