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Gametogonium (plural gametogonia) are
stem cell In multicellular organisms, stem cells are undifferentiated or partially differentiated cells that can change into various types of cells and proliferate indefinitely to produce more of the same stem cell. They are the earliest type of cell ...
s for
gamete A gamete ( ) is a Ploidy#Haploid and monoploid, haploid cell that fuses with another haploid cell during fertilization in organisms that Sexual reproduction, reproduce sexually. Gametes are an organism's reproductive cells, also referred to as s ...
s located within the
gonad A gonad, sex gland, or reproductive gland is a Heterocrine gland, mixed gland and sex organ that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gon ...
s. They originate from primordial germ cells, which have migrated to the gonads. Male gametogonia which are located within the
testes A testicle or testis ( testes) is the gonad in all male bilaterians, including humans, and is homologous to the ovary in females. Its primary functions are the production of sperm and the secretion of androgens, primarily testosterone. The ...
during development and adulthood are called
spermatogonium A spermatogonium (plural: ''spermatogonia'') is an undifferentiated male germ cell. Spermatogonia undergo spermatogenesis to form mature spermatozoa in the seminiferous tubules of the testicles. There are three subtypes of spermatogonia in human ...
(plural spermatogonia). Female gametogonia, known as
oogonium An oogonium (: oogonia) is a small diploid cell which, upon maturation, forms a primordial follicle in a female fetus or the female (haploid or diploid) gametangium of certain thallophytes. In the mammalian fetus Oogonia are formed in large ...
(plural oogonia), are found within the
ovaries The ovary () is a gonad in the female reproductive system that produces ova; when released, an ovum travels through the fallopian tube/oviduct into the uterus. There is an ovary on the left and the right side of the body. The ovaries are endocr ...
of the developing
foetus A fetus or foetus (; : fetuses, foetuses, rarely feti or foeti) is the unborn offspring of a viviparous animal that develops from an embryo. Following the embryonic stage, the fetal stage of development takes place. Prenatal development is a ...
and were thought to be depleted at or after birth. Spermatogonia and oogonia are classified as sexually differentiated germ cells.


Origin

Germ cells are specified early in development and can only differentiate into
gamete A gamete ( ) is a Ploidy#Haploid and monoploid, haploid cell that fuses with another haploid cell during fertilization in organisms that Sexual reproduction, reproduce sexually. Gametes are an organism's reproductive cells, also referred to as s ...
s. The segregation of germ cells is often determined by the species, with some undergoing preformation, where the germ cells are determined by maternally inherited factors before or immediately after fertilisation, and others undergoing epigenesis, where the germ cell lineage is determined from signalling from surrounding tissues. Preformation was initially perceived as more common than epigenesis, as it appears in many model organisms like the common fruit fly,
roundworms The nematodes ( or ; ; ), roundworms or eelworms constitute the phylum Nematoda. Species in the phylum inhabit a broad range of environments. Most species are free-living, feeding on microorganisms, but many are parasitic. Parasitic worms (hel ...
and some
amphibian Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
s. Epigenesis has since been shown to be the more common mechanism. The specific mechanism of germ line differentiation varies across species.


Animal models

Mice and other mammalian species undergo epigenesis during development, where germ cells are separated from the somatic lineage during early
gastrulation Gastrulation is the stage in the early embryonic development of most animals, during which the blastula (a single-layered hollow sphere of cells), or in mammals, the blastocyst, is reorganized into a two-layered or three-layered embryo known as ...
, occurring at embryonic day 7 in mice, and are derived directly from proximal
epiblast In amniote embryonic development, the epiblast (also known as the primitive ectoderm) is one of two distinct cell layers arising from the inner cell mass in the mammalian blastocyst, or from the blastula in reptiles and birds. It drives the em ...
cells relative to the extraembryonic
ectoderm The ectoderm is one of the three primary germ layers formed in early embryonic development. It is the outermost layer, and is superficial to the mesoderm (the middle layer) and endoderm (the innermost layer). It emerges and originates from the o ...
. Prior to
gastrulation Gastrulation is the stage in the early embryonic development of most animals, during which the blastula (a single-layered hollow sphere of cells), or in mammals, the blastocyst, is reorganized into a two-layered or three-layered embryo known as ...
the epiblast cells are not yet set in their role as cells of the germ lineage and can act as precursors for somatic cells Matsui and Okamura, 2003. At this stage, cells transplanted to the proximal epiblast from other parts of the epiblast can also be differentiated into germ line cells. The potential germ line cells are specified by the extracellular signalling of
BMP4 Bone morphogenetic protein 4 is a protein that in humans is encoded by ''BMP4'' gene. BMP4 is found on chromosome 14q22-q23. BMP4 is a member of the bone morphogenetic protein family which is part of the transforming growth factor-beta superfami ...
, BMP2 and BMP8b from the extraembryonic ectoderm. The germ cell population (~40 in mice), after specification, migrate to the developing
gonad A gonad, sex gland, or reproductive gland is a Heterocrine gland, mixed gland and sex organ that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gon ...
s where they differentiate further into gametogonia. Much of the research in germ cell development is done on
animal models A model organism is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workings of other organisms. Mod ...
. Animal models are an effective research tool due to the commonality of
sexual reproduction Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote tha ...
which is thought to have same or similar mechanisms across species. The majority of research is done on mice which has led to advances in understanding germ line differentiation across all mammal but there are some species specific mechanisms which have not been studied as extensively due to the difficulty of both obtaining human samples and the ethical limitations of human research. To circumvent that, there have been studies performed on human pluripotent stem cells.


Human research

Pluripotent stem cells are used in lieu of in vivo cells when researching germ cell development but is not without its issues. There is a limited amount of information on early germ cells, so it is difficult to ascertain if the resultant cells in the culture are the same as germ cells. Instead, research is based on inducing cells with the same or similar properties in order to study the underlying mechanisms of germ cell differentiation. Additionally, it is also difficult to compare with previous studies, especially since the majority have been done on mice, and there are different processes between species.


The role of the gonads

The gonads have an important role in germ cell development, converting the bipotential cells into gametogonia. The germ cells are bipotential in that prior to migrating to the gonads, they are capable of forming either spermatogonia or oogonia. The specification into either female or male fates for the organism itself also depends on the development of the gonads, which have yet to differentiate into ether ovaries or testes. In the mouse, somatic sex determination (i.e. determination of either female or male gonads) begins at embryonic day 10.5 in mice, but is not finalised until day 12.5. Male somatic cell specification involves the Y-chromosome specific Sry gene, which regulates the specification of
Sertoli cell Sertoli cells are a type of sustentacular "nurse" cell found in human testes which contribute to the process of spermatogenesis (the production of sperm) as a structural component of the seminiferous tubules. They are activated by follicle-sti ...
s and Leydig cells. Without the specification of those cells, the gonad formation is regulated by the X-chromosomes, forming the ovaries. After the fate of the gonads is finalised, the sex specification of the germ cells occurs at 12.5-15.5 days. Sex specification of the germ cells requires the repression of pluripotency and relies on the communication between the somatic cells of the gonads and germ cells. The mechanisms for male and female differentiation are markedly different, since a population of sperm producing spermatogonia are retained throughout development and into adult life, unlike oogonia which only produce oocyte in utero. When the germ cells reach the gonads, they undergo proliferation via mitosis and at 13.5 days of rat development they begin to undergo meiosis in the ovary but arrested at the mitotic stage in the testes. In the ovary, after
mitosis Mitosis () is a part of the cell cycle in eukaryote, eukaryotic cells in which replicated chromosomes are separated into two new Cell nucleus, nuclei. Cell division by mitosis is an equational division which gives rise to genetically identic ...
, the gametogonium undergo
meiosis Meiosis () is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, the sperm or egg cells. It involves two rounds of division that ultimately result in four cells, each with only one c ...
, which is initiated by intrinsic competence factor DazL and extrinsic
retinoic acid Retinoic acid (simplified nomenclature for all-''trans''-retinoic acid) is a metabolite of vitamin A1 (all-''trans''-retinol) that is required for embryonic development, male fertility, regulation of bone growth and immune function. All-''trans ...
, excreted by the
mesonephros The mesonephros () is one of three excretory system, excretory organs that develop in vertebrates. It serves as the main excretory organ of aquatic vertebrates and as a temporary kidney in reptiles, birds, and mammals. The mesonephros is included ...
. Retinoic acid is the major factor in meiosis, upregulating genes including ‘‘Stra8’‘, ‘‘ Dmc1’‘ and ‘‘ Sycp3’‘, which all have a role in meiosis. The male germ cells are protected from external signalling, like retinoic acid from the mesonephros, by the Leydig and Sertoli cells.


Spermatogonium

Spermatogonium are the stem cell population that reside in the testes and undergo spermatogenesis to form
sperm Sperm (: sperm or sperms) is the male reproductive Cell (biology), cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm ...
. They remain within the testes throughout adulthood.


Oogonium

Oogonium are the stem cells for
oocyte An oocyte (, oöcyte, or ovocyte) is a female gametocyte or germ cell involved in reproduction. In other words, it is an immature ovum, or egg cell. An oocyte is produced in a female fetus in the ovary during female gametogenesis. The female ger ...
s, and were previously thought to be depleted during development during the production of the primary oocytes. However, some of these stem cells have been found in the mouse and primate ovaries.


References


External links

{{cite book , last=Convery , first=Paul , title=Inkhorn's Erotonomicon: An Advanced Sexual Vocabulary for Verbivores and Vulgarians , publisher=Troubador Publishing Ltd , year=2012 , isbn=9781780883151 , url=https://books.google.com/books?id=AZfayFbCEzEC&dq=gametogonium&pg=PA96 , accessdate=8 January 2013 Germ cells