Pronucleus
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A pronucleus () is the
nucleus Nucleus ( : nuclei) is a Latin word for the seed inside a fruit. It most often refers to: *Atomic nucleus, the very dense central region of an atom *Cell nucleus, a central organelle of a eukaryotic cell, containing most of the cell's DNA Nucle ...
of a
sperm Sperm is the male reproductive 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 with a tail known as a flagellum, whi ...
or
egg cell The egg cell, or ovum (plural ova), is the female reproductive cell, or gamete, in most anisogamous organisms (organisms that reproduce sexually with a larger, female gamete and a smaller, male one). The term is used when the female gamete is ...
during the process of
fertilization Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Proce ...
. The
sperm cell Sperm is the male reproductive 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 with a tail known as a flagellum, whi ...
becomes a pronucleus after the sperm enters the ovum, but before the genetic material of the sperm and egg fuse. Contrary to the sperm cell, the egg cell has a pronucleus once it becomes haploid, and not when the sperm cell arrives. Sperm and egg cells are
haploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
, meaning they carry half the number of
chromosome A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...
s of somatic cells, so in humans, haploid cells have 23 chromosomes, while somatic cells have 46 chromosomes. The male and female pronuclei do not fuse, although their genetic material does. Instead, their membranes dissolve, leaving no barriers between the male and female chromosomes. Their chromosomes can then combine and become part of a single diploid nucleus in the resulting
embryo An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male spe ...
, containing a full set of chromosomes. The appearance of two pronuclei is the first sign of successful fertilization as observed during
in vitro fertilisation In vitro fertilisation (IVF) is a process of fertilisation where an egg is combined with sperm in vitro ("in glass"). The process involves monitoring and stimulating an individual's ovulatory process, removing an ovum or ova (egg or eggs) ...
, and is usually observed 18 hours after
insemination Insemination is the introduction of sperm into a female’s reproductive system for the purpose of impregnating, also called fertilizing, the female for sexual reproduction. The sperm is introduced into the uterus of a mammal or the oviduct of an ...
or
intracytoplasmic sperm injection Intracytoplasmic sperm injection (ICSI ) is an in vitro fertilization (IVF) procedure in which a single sperm cell is injected directly into the cytoplasm of an egg. This technique is used in order to prepare the gametes for the obtention of emb ...
(ICSI). The
zygote A zygote (, ) is a eukaryotic cell formed by a fertilization event between two gametes. The zygote's genome is a combination of the DNA in each gamete, and contains all of the genetic information of a new individual organism. In multicellula ...
is then termed a ''two-pronuclear zygote'' (2PN). Two-pronuclear zygotes transitioning through 1PN or 3PN states tend to develop into poorer-quality embryos than ones who remain 2PN throughout development, and may be significant in
embryo selection In vitro fertilisation (IVF) is a process of fertilisation where an egg is combined with sperm in vitro ("in glass"). The process involves monitoring and stimulating an individual's ovulatory process, removing an ovum or ova (egg or eggs) f ...
in
in vitro fertilisation In vitro fertilisation (IVF) is a process of fertilisation where an egg is combined with sperm in vitro ("in glass"). The process involves monitoring and stimulating an individual's ovulatory process, removing an ovum or ova (egg or eggs) ...
(IVF).


History

The pronucleus was discovered the 1870s
microscopically Microscopy is the technical field of using microscopes to view objects and areas of objects that cannot be seen with the naked eye (objects that are not within the resolution range of the normal eye). There are three well-known branches of micr ...
using staining techniques combined with
microscope A microscope () is a laboratory instrument used to examine objects that are too small to be seen by the naked eye. Microscopy is the science of investigating small objects and structures using a microscope. Microscopic means being invisibl ...
s with improved magnification levels. The pronucleus was originally found during the first studies on meiosis. Edouard Van Beneden published a paper in 1875 in which he first mentions the pronucleus by studying the eggs of rabbits and bats. He stated that the two pronuclei form together in the center of the cell to form the
embryo An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male spe ...
nic nucleus. Van Beneden also found that the sperm enters into the cell through the membrane in order to form the male pronucleus. In 1876,
Oscar Hertwig Oscar Hertwig (21 April 1849 in Friedberg – 25 October 1922 in Berlin) was a German embryologist and zoologist known for his research in developmental biology and evolution. Hertwig is credited as the first man to observe sexual reproduction ...
did a study on sea urchin eggs because the eggs of sea urchins are transparent, so it allowed for much better magnification of the egg. Hertwig confirmed Van Beneden's finding of the pronucleus, and also found the formation of the female pronucleus involves the formation of polar bodies.


Formation

The female pronucleus is the female egg cell once it has become a
haploid cell Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
, and the male pronucleus forms when the sperm enters into the female egg. While the sperm develops inside of the male
testes A testicle or testis (plural testes) is the male reproductive gland or gonad in all bilaterians, including humans. It is homologous to the female ovary. The functions of the testes are to produce both sperm and androgens, primarily testoster ...
, the sperm does not become a pronucleus until it decondenses quickly inside of the female egg.Gilbert SF. Developmental Biology. 6th edition. Sunderland (MA): Sinauer Associates; 2000. Fusion of the Genetic Material. When the sperm reaches the female egg, the sperm loses its outside membrane as well as its tail. The sperm does this because the membrane and the tail are no longer needed by the female ovum. The purpose of the cell membrane was to protect the DNA from the acidic
vaginal fluid Vaginal discharge is a mixture of liquid, cells, and bacteria that lubricate and protect the vagina. This mixture is constantly produced by the cells of the vagina and cervix, and it exits the body through the vaginal opening. The composition, amo ...
, and the purpose of the tail of the sperm was to help move the sperm cell to the egg cell. The formation of the female egg is asymmetrical, while the formation of the male sperm is symmetrical. Typically in a female mammal, meiosis starts with one diploid cell and becomes one haploid ovum and typically two
polar bodies A polar body is a small haploid cell that is formed at the same time as an egg cell during oogenesis, but generally does not have the ability to be fertilized. It is named from its polar position in the egg. When certain diploid cells in animals ...
, however one may later divide to form a third polar body. In a male, meiosis starts with one diploid cell and ends with four sperm. In mammals, the female pronucleus starts in the center of the egg before fertilization. When the male pronucleus is formed, after the sperm cell reaches the egg, the two pronuclei migrate towards each other. However, in brown alga ''
Pelvetia ''Pelvetia canaliculata'', the channelled wrack, is a very common brown alga (Phaeophyceae) found on the rocks of the upper shores of Europe. It is the only species remaining in the monotypic genus ''Pelvetia''. In 1999, the other members of this ...
'', the egg pronucleus starts in the center of the egg before fertilization and remain in the center after fertilization. This is because the egg cells of brown alga ''Pelvetia'', the egg pronucleus is anchored down by
microtubules Microtubules are polymers of tubulin that form part of the cytoskeleton and provide structure and shape to eukaryotic cells. Microtubules can be as long as 50 micrometres, as wide as 23 to 27  nm and have an inner diameter between 11 an ...
so only the male pronucleus migrates towards the female pronucleus.


Calcium concentration

The calcium concentration within the egg cell
cytoplasm In cell biology, the cytoplasm is all of the material within a eukaryotic cell, enclosed by the cell membrane, except for the cell nucleus. The material inside the nucleus and contained within the nuclear membrane is termed the nucleoplasm. The ...
has a very important role in the formation of an activated female egg. If there is no calcium influx, the female diploid cell will produce three pronuclei, rather than only one. This is due to the failure of release of the second polar body.


Combination of male and female pronuclei

In sea urchins, the formation of the
zygote A zygote (, ) is a eukaryotic cell formed by a fertilization event between two gametes. The zygote's genome is a combination of the DNA in each gamete, and contains all of the genetic information of a new individual organism. In multicellula ...
starts with the fusion of both the inner and outer nuclei of the male and female pronuclei. It is unknown if one of the pronuclei start the combination of the two, or if the microtubules that help the dissolution of membranes commence the action. The microtubules that make the two pronuclei combine come from the sperm's
centrosome In cell biology, the centrosome (Latin centrum 'center' + Greek sōma 'body') (archaically cytocentre) is an organelle that serves as the main microtubule organizing center (MTOC) of the animal cell, as well as a regulator of cell-cycle progres ...
. There is a study that strongly supports that microtubules are an important part of the fusion of the pronuclei.
Vinblastine Vinblastine (VBL), sold under the brand name Velban among others, is a chemotherapy medication, typically used with other medications, to treat a number of types of cancer. This includes Hodgkin's lymphoma, non-small cell lung cancer, bladder ca ...
is a chemotherapy drug that affects both the plus and minus ends of microtubules. When vinblastine is added to the ovum, there is a high rate of pronuclear fusion failure. This high rate of pronuclear fusion failure highly suggests that microtubules play a major role in the fusion of the pronucleus. In mammals, the pronuclei only last in the cell for about twelve hours, due to the fusion of the genetic material of the two pronuclei within the egg cell. Many studies of pronuclei have been in the egg cells of sea urchins, where the pronuclei are in the egg cell for less than an hour. The main difference between the process of fusion of genetic materials in mammals versus sea urchins is that in sea urchins, the pronuclei go directly into forming a zygote nucleus. In mammalian egg cells, the chromatin from the pronuclei form chromosomes that merge onto the same mitotic spindle. The diploid nucleus in mammals is first seen at the 2-cell stage, whereas in sea urchins it is first found at the zygote stage.


References


External links

{{wiktionary, pronucleus Genetics