![Sperm-egg](https://upload.wikimedia.org/wikipedia/commons/8/86/Sperm-egg.jpg)
Fertilisation or fertilization (see
spelling differences
Despite the various English dialects spoken from country to country and within different regions of the same country, there are only slight regional variations in English orthography, the two most notable variations being British and American ...
), also known as generative fertilisation, syngamy and impregnation, is the fusion of
gametes
A gamete (; , ultimately ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce ...
to give rise to a new individual organism or offspring and initiate its development. Processes such as
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 a ...
or
pollination
Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds, a ...
which happen before the fusion of gametes are also sometimes informally called fertilisation. The cycle of fertilisation and development of new individuals is called
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 th ...
. During
double fertilisation
Double fertilization is a complex fertilization mechanism of flowering plants (angiosperms). This process involves the joining of a female gametophyte (megagametophyte, also called the embryo sac) with two male gametes (sperm). It begins when a ...
in
angiosperms
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants ...
the
haploid male gamete combines with two haploid
polar nuclei
Double fertilization is a complex fertilization mechanism of flowering plants (angiosperms). This process involves the joining of a female gametophyte (megagametophyte, also called the embryo sac) with two male gametes (sperm). It begins when a ...
to form a
triploid
Polyploidy is a condition in which the cells of an organism have more than one pair of ( homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set contains ...
primary
endosperm nucleus by the process of vegetative fertilisation.
History
In Antiquity,
Aristotle
Aristotle (; grc-gre, Ἀριστοτέλης ''Aristotélēs'', ; 384–322 BC) was a Greek philosopher and polymath during the Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatetic school of ph ...
conceived the formation of new individuals through fusion of male and female fluids, with form and function emerging gradually, in a mode called by him as
epigenetic.
In 1784,
Spallanzani
Lazzaro Spallanzani (; 12 January 1729 – 11 February 1799) was an Italian Catholic priest (for which he was nicknamed Abbé Spallanzani), biologist and physiologist who made important contributions to the experimental study of bodily function ...
established the need of interaction between the female's ovum and male's sperm to form a zygote in frogs.
[ In 1827, von Baer observed a ]theria
Theria (; Greek: , wild beast) is a subclass of mammals amongst the Theriiformes. Theria includes the eutherians (including the placental mammals) and the metatherians (including the marsupials) but excludes the egg-laying monotremes.
...
n mammalian egg for the first time.[ ]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 ...
(1876), in Germany, described the fusion of nuclei of spermatozoa and of ova from sea urchin.
Evolution
The evolution of fertilisation is related to the origin of meiosis, as both are part 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 th ...
, originated in eukaryotes. There are two conflicting theories on how the couple meiosis–fertilisation arose. One is that it evolved from prokaryotic sex (bacterial recombination
Bacterial recombination is a type of genetic recombination in bacteria characterized by DNA transfer from one organism called donor to another organism as recipient. This process occurs in three main ways:
* Transformation, the uptake of exogeno ...
) as eukaryotes evolved from prokaryotes. The other is that mitosis originated meiosis.
Fertilisation in plants
The gametes that participate in fertilisation of plants are the sperm (male) and the egg (female) cell. Various families of plants have differing methods by which the gametes produced by the male and female gametophyte
A gametophyte () is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the ...
s come together and are fertilised. In Bryophyte
The Bryophyta s.l. are a proposed taxonomic division containing three groups of non-vascular land plants (embryophytes): the liverworts, hornworts and mosses. Bryophyta s.s. consists of the mosses only. They are characteristically limited in s ...
land plants, fertilisation of the sperm and egg takes place within the archegonium. In seed plant
A spermatophyte (; ), also known as phanerogam (taxon Phanerogamae) or phaenogam (taxon Phaenogamae), is any plant that produces seeds, hence the alternative name seed plant. Spermatophytes are a subset of the embryophytes or land plants. They inc ...
s, the male gametophyte is called a pollen grain. After pollination
Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds, a ...
, the pollen grain germinates, and a pollen tube grows and penetrates the ovule
In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: the '' integument'', forming its outer layer, the ''nucellus'' (or remnant of the megasporangium), and the ...
through a tiny pore called a micropyle Micropyle may refer to:
* Micropyle (botany)
In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: the ''integument'', forming its outer layer, the ''nucellus'' (o ...
. The sperm are transferred from the pollen through the pollen tube to the ovule where the egg is fertilised. In flowering plant
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s, two sperm cells are released from the pollen grain, and a second fertilisation event involving the second sperm cell and the central cell
Double fertilization is a complex fertilization mechanism of flowering plants (angiosperms). This process involves the joining of a female gametophyte ( megagametophyte, also called the embryo sac) with two male gametes (sperm). It begins when ...
of the ovule, which is a second female gamete.
Pollen tube growth
Unlike animal sperm which is motile, plant sperm is immotile and relies on the pollen tube to carry it to the ovule where the sperm is released. The pollen tube penetrates the stigma and elongates through the extracellular matrix of the style before reaching the ovary. Then near the receptacle, it breaks through the ovule
In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: the '' integument'', forming its outer layer, the ''nucellus'' (or remnant of the megasporangium), and the ...
through the micropyle (an opening in the ovule wall) and the pollen tube "bursts" into the embryo sac, releasing sperm. The growth of the pollen tube has been believed to depend on chemical cues from the pistil, however these mechanisms were poorly understood until 1995. Work done on tobacco plants revealed a family of glycoproteins called TTS proteins that enhanced growth of pollen tubes. Pollen tubes in a sugar free pollen germination medium and a medium with purified TTS proteins both grew. However, in the TTS medium, the tubes grew at a rate 3x that of the sugar-free medium. TTS proteins were also placed on various locations of semi in vevo pollinated pistils, and pollen tubes were observed to immediately extend toward the proteins. Transgenic plants lacking the ability to produce TTS proteins exhibited slower pollen tube growth and reduced fertility.
Rupture of pollen tube
The rupture of the pollen tube to release sperm in '' Arabidopsis'' has been shown to depend on a signal from the female gametophyte. Specific proteins called FER protein kinases present in the ovule control the production of highly reactive derivatives of oxygen called reactive oxygen species
In chemistry, reactive oxygen species (ROS) are highly reactive chemicals formed from diatomic oxygen (). Examples of ROS include peroxides, superoxide, hydroxyl radical, singlet oxygen, and alpha-oxygen.
The reduction of molecular oxygen () p ...
(ROS). ROS levels have been shown via GFP to be at their highest during floral stages when the ovule is the most receptive to pollen tubes, and lowest during times of development and following fertilisation. High amounts of ROS activate Calcium ion channels in the pollen tube, causing these channels to take up Calcium ions in large amounts. This increased uptake of calcium causes the pollen tube to rupture, and release its sperm into the ovule. Pistil feeding assays in which plants were fed diphenyl iodonium chloride (DPI) suppressed ROS concentrations in ''Arabidopsis'', which in turn prevented pollen tube rupture.
Bryophytes
Bryophyte is a traditional name used to refer to all embryophytes (land plants) that do not have true vascular tissue and are therefore called "non-vascular plants". Some bryophytes do have specialised tissues for the transport of water; however, since these do not contain lignin, they are not considered true vascular tissue.
Ferns
A fern is a member of a group of roughly 12,000 species of vascular plants that reproduce via spores and have neither seeds nor flowers. They differ from mosses by being vascular (i.e. having water-conducting vessels). They have stems and leaves, like other vascular plants. Most ferns have what are called fiddleheads that expand into fronds, which are each delicately divided.
Gymnosperms
The gymnosperms are a group of seed producing plants that includes conifers, Cycad
Cycads are seed plants that typically have a stout and woody (ligneous) trunk with a crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants of a species are either male o ...
s, Ginkgo
''Ginkgo'' is a genus of non-flowering seed plants. The scientific name is also used as the English name. The order to which it belongs, Ginkgoales, first appeared in the Permian, 270 million years ago, and is now the only living genus with ...
, and Gnetales
Gnetophyta () is a division of plants (alternatively considered the subclass Gnetidae or order Gnetales), grouped within the gymnosperms (which also includes conifers, cycads, and ginkgos), that consists of some 70 species across the three reli ...
. The term "gymnosperm" comes from the Greek composite word γυμνόσπερμος (γυμνός gymnos, "naked" and σπέρμα sperma, "seed"), meaning "naked seeds", after the unenclosed condition of their seeds (called ovules in their unfertilised state). Their naked condition stands in contrast to the seeds and ovules of flowering plants (angiosperms), which are enclosed within an ovary. Gymnosperm seeds develop either on the surface of scales or leaves, often modified to form cones, or at the end of short stalks as in Ginkgo.
Flowering plants
After being fertilised, the ovary starts to swell and develop into the fruit
In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering.
Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in particu ...
. With multi-seeded fruits, multiple grains of pollen are necessary for syngamy with each ovule. The growth of the pollen tube is controlled by the vegetative (or tube) cytoplasm. Hydrolytic enzymes are secreted by the pollen tube that digest the female tissue as the tube grows down the stigma and style; the digested tissue is used as a nutrient source for the pollen tube as it grows. During pollen tube growth towards the ovary, the generative nucleus divides to produce two separate sperm nuclei (haploid number of chromosomes) – a growing pollen tube therefore contains three separate nuclei, two sperm and one tube. The sperms are interconnected and dimorphic, the large one, in a number of plants, is also linked to the tube nucleus and the interconnected sperm and the tube nucleus form the "male germ unit".
Double fertilisation
Double fertilization is a complex fertilization mechanism of flowering plants (angiosperms). This process involves the joining of a female gametophyte (megagametophyte, also called the embryo sac) with two male gametes (sperm). It begins when a ...
is the process in angiosperms
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants ...
(flowering plants) in which two sperm from each pollen tube fertilise two cells in a female gametophyte
A gametophyte () is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the ...
(sometimes called an embryo sac) that is inside an ovule. After the pollen tube enters the gametophyte, the pollen tube nucleus disintegrates and the two sperm cells are released; one of the two sperm cells ''fertilises'' the egg cell (at the bottom of the gametophyte near the micropyle), forming a diploid (2n) 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 multicell ...
. This is the point when fertilisation actually occurs; pollination and fertilisation are two separate processes. The nucleus of the other sperm cell fuses with two haploid polar nuclei (contained in the central cell) in the centre of the gametophyte. The resulting cell is triploid
Polyploidy is a condition in which the cells of an organism have more than one pair of ( homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set contains ...
(3n). This triploid cell divides through mitosis and forms the endosperm, a nutrient-rich tissue, inside the seed
A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiospe ...
.
The two central-cell maternal nuclei (polar nuclei) that contribute to the endosperm arise by mitosis from the single meiotic product that also gave rise to the egg. Therefore, maternal contribution to the genetic constitution of the triploid endosperm is double that of the embryo.
One primitive species of flowering plant, ''Nuphar polysepala
''Nuphar polysepala'' (the great yellow pond-lily or wokas; syn. ''Nuphar polysepalum'' orth. err., ''Nuphar lutea'' subsp. ''polysepala'' (Engelm.) E.O.Beal) is a species of '' Nuphar'' native to western North America.Flora of North America''Nup ...
'', has endosperm that is diploid, resulting from the fusion of a sperm with one, rather than two, maternal nuclei. It is believed that early in the development of angiosperm linages, there was a duplication in this mode of reproduction, producing seven-celled/eight-nucleate female gametophytes, and triploid endosperms with a 2:1 maternal to paternal genome ratio.
In many plants, the development of the flesh of the fruit is proportional to the percentage of fertilised ovules. For example, with watermelon
Watermelon (''Citrullus lanatus'') is a flowering plant species of the Cucurbitaceae family and the name of its edible fruit. A scrambling and trailing vine-like plant, it is a highly cultivated fruit worldwide, with more than 1,000 varie ...
, about a thousand grains of pollen must be delivered and spread evenly on the three lobes of the stigma to make a normal sized and shaped fruit.
Cross-fertilisation and self-fertilisation represent different strategies with differing benefits and costs. An estimated 48.7% of plant species are either dioecious or self-incompatible obligate out-crossers. It is also estimated that about 42% of flowering plants exhibit a mixed mating system in nature.
In the most common kind of mixed mating system, individual plants produce a single type of flower and fruits may contain self-fertilised, out-crossed or a mixture of progeny types. The transition from cross-fertilisation to self-fertilisation is the most common evolutionary transition in plants, and has occurred repeatedly in many independent lineages. About 10-15% of flowering plants are predominantly self-fertilising.
Self-pollination
Under circumstances where pollinators or mates are rare, self-fertilisation offers the advantage of reproductive assurance. Self-fertilisation can therefore result in improved colonisation ability. In some species, self-fertilisation has persisted over many generations. ''Capsella rubella
''Capsella rubella'', the pink shepherd's-purse, is a plant species in the genus '' Capsella'', a very close relative of ''Arabidopsis thaliana'' and a member of the mustard family, Brassicaceae. It has a very similar appearance to ''Capsella bur ...
'' is a self-fertilisating species that became self-compatible 50,000 to 100,000 years ago. '' Arabidopsis thaliana'' is a predominantly self-fertilising plant with an out-crossing
Out-crossing or out-breeding is the technique of crossing between different breeds. This is the practice of introducing distantly related genetic material into a breeding line, thereby increasing genetic diversity.
Outcrossing can be a usefu ...
rate in the wild of less than 0.3%; a study suggested that self-fertilisation evolved roughly a million years ago or more in ''A. thaliana''. In long-established self-fertilising plants, the masking of deleterious mutation
In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA replication, DNA or viral repl ...
s and the production of genetic variability is infrequent and thus unlikely to provide a sufficient benefit over many generations to maintain the meiotic apparatus. Consequently, one might expect self-fertilisation to be replaced in nature by an ameiotic asexual form of reproduction that would be less costly. However the actual persistence of meiosis and self-fertilisation as a form of reproduction in long-established self-fertilising plants may be related to the immediate benefit of efficient recombinational repair
Homologous recombination is a type of genetic recombination in which genetic information is exchanged between two similar or identical molecules of double-stranded or single-stranded nucleic acids (usually DNA as in cellular organisms but may be ...
of DNA damage during formation of germ cells provided by meiosis at each generation.
Fertilisation in animals
The mechanics behind fertilisation has been studied extensively in sea urchins and mice. This research addresses the question of how the sperm and the appropriate egg find each other and the question of how only one sperm gets into the egg and delivers its contents. There are three steps to fertilisation that ensure species-specificity:
# Chemotaxis
# Sperm activation/acrosomal reaction
# Sperm/egg adhesion
Internal vs. external
Consideration as to whether an animal (more specifically a vertebrate) uses internal
Internal may refer to:
*Internality as a concept in behavioural economics
*Neijia, internal styles of Chinese martial arts
*Neigong or "internal skills", a type of exercise in meditation associated with Daoism
*''Internal (album)'' by Safia, 2016
...
or external fertilisation
External fertilization is a mode of reproduction in which a male organism's sperm fertilizes a female organism's egg outside of the female's body.
It is contrasted with internal fertilization, in which sperm are introduced via insemination and then ...
is often dependent on the method of birth. Oviparous animals laying eggs with thick calcium shells, such as chickens, or thick leathery shells generally reproduce via internal fertilisation so that the sperm fertilises the egg without having to pass through the thick, protective, tertiary layer of the egg. Ovoviviparous
Ovoviviparity, ovovivipary, ovivipary, or aplacental viviparity is a term used as a "bridging" form of reproduction between egg-laying oviparous and live-bearing viviparous reproduction. Ovoviviparous animals possess embryos that develop insi ...
and viviparous
Among animals, viviparity is development of the embryo inside the body of the parent. This is opposed to oviparity which is a reproductive mode in which females lay developing eggs that complete their development and hatch externally from the ...
animals also use internal fertilisation. It is important to note that although some organisms reproduce via amplexus
Amplexus (Latin "embrace") is a type of mating behavior exhibited by some externally fertilizing species (chiefly amphibians and horseshoe crabs) in which a male grasps a female with his front legs as part of the mating process, and at the same ...
, they may still use internal fertilisation, as with some salamanders. Advantages to internal fertilisation include: minimal waste of gametes; greater chance of individual egg fertilisation, relatively "longer" time period of egg protection, and selective fertilisation; many females have the ability to store sperm for extended periods of time and can fertilise their eggs at their own desire.
Oviparous animals producing eggs with thin tertiary membranes or no membranes at all, on the other hand, use external fertilisation methods. Such animals may be more precisely termed ovuliparous. Advantages to external fertilisation include: minimal contact and transmission of bodily fluids; decreasing the risk of disease transmission, and greater genetic variation (especially during broadcast spawning external fertilisation methods).
Sea urchins
Sperm find the eggs via chemotaxis, a type of ligand/receptor interaction. Resact is a 14 amino acid peptide purified from the jelly coat of ''A. punctulata'' that attracts the migration of sperm.
After finding the egg, the sperm penetrates the jelly coat through a process called sperm activation. In another ligand/receptor interaction, an oligosaccharide component of the egg binds and activates a receptor on the sperm and causes the acrosomal reaction
During fertilization, a sperm must first fuse with the plasma membrane and then penetrate the female egg cell to fertilize it. Fusing to the egg cell usually causes little problem, whereas penetrating through the egg's hard shell or extracellular ...
. The acrosomal vesicles of the sperm fuse with the plasma membrane and are released. In this process, molecules bound to the acrosomal vesicle membrane, such as bindin, are exposed on the surface of the sperm. These contents digest the jelly coat and eventually the vitelline membrane. In addition to the release of acrosomal vesicles, there is explosive polymerisation of actin
Actin is a family of globular multi-functional proteins that form microfilaments in the cytoskeleton, and the thin filaments in muscle fibrils. It is found in essentially all eukaryotic cells, where it may be present at a concentration of ov ...
to form a thin spike at the head of the sperm called the acrosomal process The acrosome is an organelle that develops over the anterior (front) half of the head in the spermatozoa (sperm cells) of many animals including humans. It is a cap-like structure derived from the Golgi apparatus. In placental mammals the acrosome c ...
.
The sperm binds to the egg through another ligand reaction between receptors on the vitelline membrane
The vitelline membrane or vitelline envelope is a structure surrounding the outer surface of the plasma membrane of an ovum (the oolemma) or, in some animals (e.g., birds), the extracellular yolk and the oolemma. It is composed mostly of protein f ...
. The sperm surface protein bindin, binds to a receptor on the vitelline membrane identified as EBR1.
Fusion of the plasma membranes of the sperm and egg are likely mediated by bindin. At the site of contact, fusion causes the formation of a fertilisation cone.
Mammals
Mammals internally fertilise through copulation
Sexual intercourse (or coitus or copulation) is a sexual activity typically involving the insertion and thrusting of the penis into the vagina for sexual pleasure or reproduction.Sexual intercourse most commonly means penile–vaginal penetra ...
. After a male ejaculates
Ejaculation is the discharge of semen (the ''ejaculate''; normally containing sperm) from the male reproductory tract as a result of an orgasm. It is the final stage and natural objective of male sexual stimulation, and an essential componen ...
, many sperm move to the upper vagina (via contractions from the vagina) through the cervix and across the length of the uterus
The uterus (from Latin ''uterus'', plural ''uteri'') or womb () is the organ in the reproductive system of most female mammals, including humans that accommodates the embryonic and fetal development of one or more embryos until birth. The ...
to meet the ovum. In cases where fertilisation occurs, the female usually ovulates during a period that extends from hours before copulation to a few days after; therefore, in most mammals it is more common for ejaculation to precede ovulation than vice versa.
When sperm are deposited into the anterior vagina, they are not capable of fertilisation (i.e., non-capacitated) and are characterized by slow linear motility patterns. This motility, combined with muscular contractions enables sperm transport towards the uterus and fallopian tubes. There is a pH gradient within the micro-environment of the female reproductive tract such that the pH near the vaginal opening is lower (approximately 5) than the fallopian tubes (approximately 8). The sperm-specific pH-sensitive calcium transport protein called CatSper increases the sperm cell permeability to calcium as it moves further into the reproductive tract. Intracellular calcium influx contributes to sperm capacitation and hyperactivation, causing a more violent and rapid non-linear motility pattern as sperm approach the oocyte. The capacitated spermatozoon and the oocyte meet and interact in the ''ampulla'' of the fallopian tube. Rheotaxis, thermotaxis and chemotaxis are known mechanisms that guide sperm towards the egg during the final stage of sperm migration. Spermatozoa respond (see Sperm thermotaxis) to the temperature gradient of ~2 °C between the oviduct and the ampulla, and chemotactic
Chemotaxis (from '' chemo-'' + '' taxis'') is the movement of an organism or entity in response to a chemical stimulus. Somatic cells, bacteria, and other single-cell or multicellular organisms direct their movements according to certain chemica ...
gradients of progesterone have been confirmed as the signal emanating from the cumulus oophorus
The cumulus oophorus, (discus proligerus), is a cluster of cells that surround the oocyte both in the ovarian follicle and after ovulation. In the antral follicle, it may be regarded as an extension of the membrana granulosa. The innermost layer ...
cells surrounding rabbit and human oocytes. Capacitated and hyperactivated sperm respond to these gradients by changing their behaviour and moving towards the cumulus-oocyte complex. Other chemotactic signals such as formyl Met-Leu-Phe (fMLF) may also guide spermatozoa.
The zona pellucida
The zona pellucida (plural zonae pellucidae, also egg coat or pellucid zone) is a specialized extracellular matrix that surrounds the plasma membrane of mammalian oocytes. It is a vital constitutive part of the oocyte. The zona pellucida first ap ...
, a thick layer of extracellular matrix that surrounds the egg and is similar to the role of the vitelline membrane in sea urchins, binds the sperm. Unlike sea urchins, the sperm binds to the egg before the acrosomal reaction. ZP3
Zona pellucida sperm-binding protein 3, also known as zona pellucida glycoprotein 3 (Zp-3) or the sperm receptor, is a ZP module-containing protein that in humans is encoded by the ''ZP3'' gene. ZP3 is the receptor in the zona pellucida which bin ...
, a glycoprotein in the zona pellucida, is responsible for egg/sperm adhesion in humans. The receptor galactosyltransferase
Galactosyltransferase is a type of glycosyltransferase which catalyzes the transfer of galactose. An example is B-N-acetylglucosaminyl-glycopeptide b-1,4-galactosyltransferase.
The biosynthesis of disaccharides, oligosaccharides and polysacchar ...
(GalT) binds to the N-acetylglucosamine residues on the ZP3 and is important for binding with the sperm and activating the acrosome reaction. ZP3 is sufficient though unnecessary for sperm/egg binding. Two additional sperm receptors exist: a 250kD protein that binds to an oviduct secreted protein, and SED1, which independently binds to the zona. After the acrosome reaction, the sperm is believed to remain bound to the zona pellucida through exposed ZP2 receptors. These receptors are unknown in mice but have been identified in guinea pigs.
In mammals, the binding of the spermatozoon to the GalT initiates the acrosome reaction
During fertilization, a sperm must first fuse with the plasma membrane and then penetrate the female egg cell to fertilize it. Fusing to the egg cell usually causes little problem, whereas penetrating through the egg's hard shell or extracellular ...
. This process releases the hyaluronidase
Hyaluronidases are a family of enzymes that catalyse the degradation of hyaluronic acid (HA). Karl Meyer classified these enzymes in 1971, into three distinct groups, a scheme based on the enzyme reaction products. The three main types of hyal ...
that digests the matrix of hyaluronic acid in the vestments around the oocyte. Additionally, heparin-like glycosaminoglycans (GAGs) are released near the oocyte that promote the acrosome reaction. Fusion between the oocyte plasma membranes and sperm follows and allows the sperm 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
Nucl ...
, the typical centriole, and atypical centriole that is attached to the flagellum
A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates.
A microorganism may have f ...
, but not the mitochondria, to enter the oocyte. The protein CD9
CD9 is a gene encoding a protein that is a member of the transmembrane 4 superfamily also known as the tetraspanin family. It is a cell surface glycoprotein that consists of four transmembrane regions and has two extracellular loops that contain ...
likely mediates this fusion in mice (the binding homolog). The egg " activates" itself upon fusing with a single sperm cell and thereby changes its cell membrane to prevent fusion with other sperm. Zinc
Zinc is a chemical element with the symbol Zn and atomic number 30. Zinc is a slightly brittle metal at room temperature and has a shiny-greyish appearance when oxidation is removed. It is the first element in group 12 (IIB) of the periodi ...
atoms are released during this activation.
This process ultimately leads to the formation of a diploid cell called a 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 multicell ...
. The zygote divides to form a blastocyst and, upon entering the uterus, implants in the endometrium, beginning pregnancy
Pregnancy is the time during which one or more offspring develops (gestation, gestates) inside a woman, woman's uterus (womb). A multiple birth, multiple pregnancy involves more than one offspring, such as with twins.
Pregnancy usually occur ...
. Embryonic implantation not in the uterine
The uterus (from Latin ''uterus'', plural ''uteri'') or womb () is the organ in the reproductive system of most female mammals, including humans that accommodates the embryonic and fetal development of one or more embryos until birth. The ...
wall results in an ectopic pregnancy that can kill the mother.
In such animals as rabbits, coitus induces ovulation by stimulating the release of the pituitary hormone gonadotropin; this release greatly increases the likelihood of pregnancy.
Humans
Fertilisation in humans is the union of a human egg
An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the a ...
and sperm, usually occurring in the ampulla of the fallopian tube, producing a 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 multicell ...
, or fertilised egg, initiating embryonic development. Scientists discovered the dynamics of human fertilisation in the nineteenth century.
The term ''conception'' commonly refers to "the process of becoming pregnant involving fertilization or implantation or both". Its use makes it a subject of semantic arguments about the beginning of pregnancy, typically in the context of the abortion
Abortion is the termination of a pregnancy by removal or expulsion of an embryo or fetus. An abortion that occurs without intervention is known as a miscarriage or "spontaneous abortion"; these occur in approximately 30% to 40% of pre ...
debate.
Upon gastrulation, which occurs around 16 days after fertilisation, the implanted blastocyst develops three germ layers, the endoderm, the ectoderm and the mesoderm, and the genetic code of the father becomes fully involved in the development of the embryo; later twinning is impossible. Additionally, interspecies hybrids survive only until gastrulation and cannot further develop.
However, some human developmental biology literature refers to the ''conceptus
A conceptus (from Latin: ''concipere'' to conceive) is an embryo and its appendages (adnexa), the associated membranes, placenta, and umbilical cord; the products of conception or, more broadly, "the product of conception at any point between fert ...
'' and such medical literature refers to the "products of conception" as the post-implantation embryo and its surrounding membranes. The term "conception" is not usually used in scientific literature because of its variable definition and connotation.
Insects
Insects in different groups, including the Odonata
Odonata is an order of flying insects that includes the dragonflies and damselflies. Members of the group first appeared during the Triassic, though members of their total group, Odonatoptera, first appeared in Late Carboniferous.
The two com ...
(dragonflies
A dragonfly is a flying insect belonging to the infraorder Anisoptera below the order Odonata. About 3,000 extant species of true dragonfly are known. Most are tropical, with fewer species in temperate regions. Loss of wetland habitat threa ...
and damselflies) and the Hymenoptera (ant
Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of ...
s, bees, and wasp
A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. ...
s) practise delayed fertilisation. Among the Odonata, females may mate with multiple males, and store sperm until the eggs are laid. The male may hover above the female during egg-laying (oviposition) to prevent her from mating with other males and replacing his sperm; in some groups such as the darters, the male continues to grasp the female with his claspers during egg-laying, the pair flying around in tandem. Among social Hymenoptera, honeybee
A honey bee (also spelled honeybee) is a eusocial flying insect within the genus ''Apis'' of the bee clade, all native to Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the current cosm ...
queens mate only on mating flights, in a short period lasting some days; a queen may mate with eight or more drones. She then stores the sperm for the rest of her life, perhaps for five years or more.
Fertilisation in fungi
In many fungi
A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...
(except chytrids), as in some protists, fertilisation is a two step process. First, the cytoplasms of the two gamete cells fuse (called plasmogamy
Plasmogamy is a stage in the sexual reproduction of fungi, in which the protoplasm of two parent cells (usually from the mycelia) fuse without the fusion of nuclei, effectively bringing two haploid nuclei close together in the same cell. This sta ...
), producing a dikaryotic The dikaryon is a cell nucleus, nuclear feature which is unique to certain fungi. (The green alga ''Derbesia'' had been long considered an exception, until the heterokaryotic hypothesis was challenged by later studies.) Compatible cell-types can fus ...
or heterokaryotic
A heterokaryon is a multinucleate cell that contains genetically different nuclei. Heterokaryotic and heterokaryosis are derived terms. This is a special type of syncytium. This can occur naturally, such as in the mycelium of fungi during sexual ...
cell with multiple nuclei. This cell may then divide to produce dikaryotic or heterokaryotic hyphae. The second step of fertilisation is karyogamy
Karyogamy is the final step in the process of fusing together two haploid eukaryotic cells, and refers specifically to the fusion of the two nuclei. Before karyogamy, each haploid cell has one complete copy of the organism's genome. In order for ...
, the fusion of the nuclei to form a diploid zygote.
In chytrid fungi, fertilisation occurs in a single step with the fusion of gametes, as in animals and plants.
Fertilisation in protists
Fertilisation in protozoa
There are three types of fertilisation processes in protozoa:
* gametogamy;
* autogamy;
* gamontogamy.
Fertilisation in algae
Fertilisation in algae occurs by binary fission
Binary may refer to:
Science and technology Mathematics
* Binary number, a representation of numbers using only two digits (0 and 1)
* Binary function, a function that takes two arguments
* Binary operation, a mathematical operation that ta ...
. The pseudopodia is first withdrawn and the nucleus starts dividing. When the cytoplasm is divided, the cytoplasm is also divided into two equal parts for each daughter cell. Two daughter cells are produced by one parent cell. It involves the process of mitosis.
Algae, like some land plants, undergo alternation of generations. Some algae are isomorphic, where both the sporophyte (2n) and gameteophyte (n) are the same morphologically. When algae reproduction is described as oogamous, the male and female gametes are different morphologically, where there is a large non-motile egg for female gametes, and the male gamete are uniflagellate (motile). Via the process of syngamy, these will form a new zygote, regenerating the sporophyte generation again.
Fertilisation in fungi-like protists
Fertilisation in fungi. In many fungi (except chytrids), as in some protists, fertilisation is a two step process. ... In chytrid fungi, fertilisation occurs in a single step with the fusion of gametes, as in animals and plants.
Fertilisation and genetic recombination
Meiosis
Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately r ...
results in a random segregation of the genes that each parent contributes. Each parent organism is usually identical save for a fraction of their genes; each gamete
A gamete (; , ultimately ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce ...
is therefore genetically unique. At fertilisation, parental 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 combine. In human
Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, cultu ...
s, (2²²)² = 17.6x1012 chromosomally different 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 multicell ...
s are possible for the non-sex chromosomes, even assuming no chromosomal crossover. If crossover occurs once, then on average (4²²)² = 309x1024 genetically different zygotes are possible for every couple, not considering that crossover events can take place at most points along each chromosome. The X and Y chromosomes undergo no crossover events and are therefore excluded from the calculation. The mitochondrial DNA is only inherited from the maternal parent.
The sperm aster and zygote centrosomes
Shortly after the sperm fuse with the egg, the two sperm centrioles form the embryo first 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 prog ...
and microtubule aster
Aster or ASTER may refer to:
Biology
* ''Aster'' (genus), a genus of flowering plants
** List of ''Aster'' synonyms, other genera formerly included in ''Aster'' and still called asters in English
* Aster (cell biology), a cellular structure shap ...
. The sperm centriole, found near the male pronucleus, recruit egg Pericentriolar material Pericentriolar material (PCM, sometimes also called pericent matrix) is a highly structured, dense mass of protein which makes up the part of the animal centrosome that surrounds the two centrioles. The PCM contains proteins responsible for microtub ...
proteins forming the zygote first centrosome. This centrosome nucleates microtubules in the shape of stars called astral microtubules. The microtubules span the whole valium of the egg, allowing the egg pronucleus to use the cables to get to the male pronucleus. As the male and female pronuclei approach each other, the single centrosome split into two centrosomes located in the interphase between the pronuclei. Then the centrosome via the astral microtubules polarizes the genome inside the pronuclei.
Parthenogenesis
Organisms that normally reproduce sexually can also reproduce via parthenogenesis
Parthenogenesis (; from the Greek grc, παρθένος, translit=parthénos, lit=virgin, label=none + grc, γένεσις, translit=génesis, lit=creation, label=none) is a natural form of asexual reproduction in which growth and developmen ...
, wherein an unfertilised female gamete produces viable offspring. These offspring may be clones of the mother, or in some cases genetically differ from her but inherit only part of her DNA. Parthenogenesis occurs in many plants and animals and may be induced in others through a chemical or electrical stimulus to the egg cell. In 2004, Japanese researchers led by Tomohiro Kono
Tomohiro is a masculine Japanese given name.
Possible writings
Tomohiro can be written using many different combinations of kanji characters. Some examples:
*友弘, "friend, vast"
*友広, "friend, wide"
*友寛, "friend, generosity"
*友博, ...
succeeded after 457 attempts to merge the ova
, abbreviated as OVA and sometimes as OAV (original animation video), are Japanese animated films and series made specially for release in home video formats without prior showings on television or in theaters, though the first part of an OVA s ...
of two mice by blocking certain proteins that would normally prevent the possibility; the resulting embryo normally developed into a mouse.
Allogamy and autogamy
Allogamy Allogamy or cross-fertilization is the fertilization of an ovum from one individual with the spermatozoa of another. By contrast, autogamy is the term used for self-fertilization. In humans, the fertilization event is an instance of allogamy. Self-f ...
, which is also known as cross-fertilisation, refers to the fertilisation of an egg cell from one individual with the male gamete of another.
Autogamy which is also known as self-fertilisation, occurs in such hermaphroditic organisms as plants and flatworms; therein, two gametes from one individual fuse.
Other variants of bisexual reproduction
Some relatively unusual forms of reproduction are:
Gynogenesis Gynogenesis, a form of parthenogenesis, is a system of asexual reproduction that requires the presence of sperm without the actual contribution of its DNA for completion. The paternal DNA dissolves or is destroyed before it can fuse with the egg. Th ...
: A sperm stimulates the egg to develop without fertilisation or syngamy. The sperm may enter the egg.
Hybridogenesis
Parthenogenesis (; from the Greek grc, παρθένος, translit=parthénos, lit=virgin, label=none + grc, γένεσις, translit=génesis, lit=creation, label=none) is a natural form of asexual reproduction in which growth and development ...
: One genome is eliminated to produce haploid eggs.
Canina meiosis
''Rosa canina'', commonly known as the dog rose, is a variable climbing, wild rose species native to Europe, northwest Africa, and western Asia.
Description
The dog rose is a deciduous shrub normally ranging in height from , though sometimes it ...
: (sometimes called "permanent odd polyploidy") one genome is transmitted in the Mendelian fashion, others are transmitted clonally.
Benefits of cross-fertilisation
The major benefit of cross-fertilisation is generally thought to be the avoidance of inbreeding depression
Inbreeding depression is the reduced biological fitness which has the potential to result from inbreeding (the breeding of related individuals). Biological fitness refers to an organism's ability to survive and perpetuate its genetic material. ...
. Charles Darwin
Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended ...
, in his 1876 book '' The Effects of Cross and Self Fertilisation in the Vegetable Kingdom'' (pages 466-467) summed up his findings in the following way.
"It has been shown in the present volume that the offspring from the union of two distinct individuals, especially if their progenitors have been subjected to very different conditions, have an immense advantage in height, weight, constitutional vigour and fertility over the self-fertilised offspring from one of the same parents. And this fact is amply sufficient to account for the development of the sexual elements, that is, for the genesis of the two sexes."
In addition, it is thought by some, that a long-term advantage of out-crossing in nature is increased genetic variability that promotes adaptation or avoidance of extinction (see Genetic variability
Genetic variability is either the presence of, or the generation of, genetic differences.
It is defined as "the formation of individuals differing in genotype, or the presence of genotypically different individuals, in contrast to environmentally i ...
).
See also
* Cell fusion
* Conception cap
A conception device is a medical device which is used to assist in the achievement of a pregnancy, often, but not always, by means other than sexual intercourse (natural insemination, or NI). This article deals exclusively with conception devices ...
* Conception device
* Female sperm Female sperm can refer to either:
#A sperm which contains an X chromosome, produced in the usual way in the testicles, referring to the occurrence of such a sperm fertilizing an egg and giving birth to a female.
#A sperm which artificially contains ...
* Fetal development
Prenatal development () includes the development of the embryo and of the fetus during a viviparous animal's gestation. Prenatal development starts with fertilization, in the germinal stage of embryonic development, and continues in fetal devel ...
* 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) ...
* Kaguya (mouse)
Kaguya was a mouse that had two parents of the same sex (). She was named after a Japanese folk tale, in which the Moon-born princess Kaguya (Kaguya-hime) is found as a baby inside a bamboo stalk.
The process of parthenogenesis
A process calle ...
* Parthenogenesis
Parthenogenesis (; from the Greek grc, παρθένος, translit=parthénos, lit=virgin, label=none + grc, γένεσις, translit=génesis, lit=creation, label=none) is a natural form of asexual reproduction in which growth and developmen ...
, a type of reproduction that does not involve fertilisation
* Pollination
Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds, a ...
* Pre-embryo
In human embryonic development a pre-embryo is a conceptus before implantation in the uterus.
Pre-embryo in human embryonic development
The word pre-embryo sometimes is used in ethical contexts to refer to a human conceptus at least between fer ...
* Pronucleus
A pronucleus () is the nucleus of a sperm or egg cell during the process of fertilization. The sperm cell 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, ...
* Superfecundation
Superfecundation is the fertilization of two or more ova from the same cycle by sperm from separate acts of sexual intercourse, which can lead to twin babies from two separate biological fathers. The term superfecundation is derived from fecund ...
* Superfetation
Superfetation (also spelled superfoetation – see fetus) is the simultaneous occurrence of more than one stage of developing offspring in the same animal.
In mammals, it manifests as the formation of an embryo from a different menstrual cycle w ...
* Symmetry breaking and cortical rotation
Symmetry breaking in biology is the process by which uniformity is broken, or the number of points to view invariance are reduced, to generate a more structured and improbable state. That is to say, symmetry breaking is the event where symmetry ...
References
External links
Fertilisation (Conception)
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Reproduction
Fertility
Pollination