Double fertilization or double fertilisation (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 Americ ...

) is a complex

fertilization Fertilisation or fertilization (see American and British English spelling differences#-ise, -ize (-isation, -ization), spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give ...

mechanism of

angiosperms Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed within a fruit. T ...

. This process involves the fusion of 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 se ...

or megagametophyte, also called the embryonic sac, with two male

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 (sperm). It begins when a pollen grain adheres to the stigmatic surface of the

carpel Gynoecium (; ; : gynoecia) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more ...

, the female reproductive structure of angiosperm flowers. The pollen grain begins to

germinate Germination is the process by which an organism grows from a seed or spore. The term is applied to the sprouting of a seedling from a seed of an flowering plant, angiosperm or gymnosperm, the growth of a sporeling from a spore, such as the sp ...

(unless a type of self-incompatibility that acts in the stigma occurs in that particular species and is activated), forming a pollen tube that penetrates and extends down through the style toward the

ovary 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 end ...

as it follows chemical signals released by the egg. The tip of the pollen tube then enters the ovary by penetrating through the micropyle opening in 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 sporangium, megasporangium), ...

, and releases two sperm into the embryonic sac (megagametophyte). The mature embryonic sac of an unfertilized ovule is 7-cellular and 8-nucleate. It is arranged in the form of 3+1+3 (from top to bottom) i.e. 3 antipodal cells, 1 central cell (binucleate), 2 synergids & 1 egg cell. One sperm fertilizes the egg cell and the other sperm fuses with the two polar nuclei of the large central cell of the megagametophyte. The

haploid Ploidy () is the number of complete sets of chromosomes in a cell (biology), cell, and hence the number of possible alleles for Autosome, autosomal and Pseudoautosomal region, pseudoautosomal genes. Here ''sets of chromosomes'' refers to the num ...

sperm and haploid egg fuse to form a

diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Here ''sets of chromosomes'' refers to the number of maternal and paternal chromosome copies, ...

zygote, the process being called syngamy, while the other sperm and the

central cell fuse to form a triploid primary endosperm cell (triple fusion). Some plants may form

polyploid Polyploidy is a condition in which the biological cell, cells of an organism have more than two paired sets of (Homologous chromosome, homologous) chromosomes. Most species whose cells have Cell nucleus, nuclei (eukaryotes) are diploid, meaning ...

nuclei. The large cell of the gametophyte will then develop into the

endosperm The endosperm is a tissue produced inside the seeds of most of the flowering plants following double fertilization. It is triploid (meaning three chromosome sets per nucleus) in most species, which may be auxin-driven. It surrounds the Embryo#Pla ...

, a nutrient-rich tissue which nourishes the developing embryo. The

, surrounding the ovules, develops into the fruit, which protects the seeds and may function to disperse them. The two central cell maternal nuclei (polar nuclei) that contribute to the endosperm, arise by mitosis from the same single meiotic product that gave rise to the egg. The maternal contribution to the genetic constitution of the triploid endosperm is double that of the sperm. In a study conducted in 2008 of the plant ''

Arabidopsis thaliana ''Arabidopsis thaliana'', the thale cress, mouse-ear cress or arabidopsis, is a small plant from the mustard family (Brassicaceae), native to Eurasia and Africa. Commonly found along the shoulders of roads and in disturbed land, it is generally ...

'', the migration of male nuclei inside the female gamete, in fusion with the female nuclei, has been documented for the first time using

in vivo Studies that are ''in vivo'' (Latin for "within the living"; often not italicized in English) are those in which the effects of various biological entities are tested on whole, living organisms or cells, usually animals, including humans, an ...

imaging. Some of the genes involved in the migration and fusion process have also been determined. Evidence of double fertilization in

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 Relict ...

, which are non-flowering seed plants, has been reported.

Brief history

Double fertilization was discovered more than a century ago by Sergei Nawaschin in

Kyiv Kyiv, also Kiev, is the capital and most populous List of cities in Ukraine, city of Ukraine. Located in the north-central part of the country, it straddles both sides of the Dnieper, Dnieper River. As of 1 January 2022, its population was 2, ...

, and Léon Guignard in

France France, officially the French Republic, is a country located primarily in Western Europe. Overseas France, Its overseas regions and territories include French Guiana in South America, Saint Pierre and Miquelon in the Atlantic Ocean#North Atlan ...

. Each made the discovery independently of the other. '' Lilium martagon'' and '' Fritillaria tenella'' were used in the first observations of double fertilization, which were made using the classical

light microscope The optical microscope, also referred to as a light microscope, is a type of microscope that commonly uses visible spectrum, visible light and a system of lens (optics), lenses to generate magnified images of small objects. Optical microscopes ...

. Due to the limitations of the light microscope, there were many unanswered questions regarding the process of double fertilization. However, with the development of the

electron microscope An electron microscope is a microscope that uses a beam of electrons as a source of illumination. It uses electron optics that are analogous to the glass lenses of an optical light microscope to control the electron beam, for instance focusing it ...

, many of the questions were answered. Most notably, the observations made by the group of W. Jensen showed that the male gametes did not have any

cell wall A cell wall is a structural layer that surrounds some Cell type, cell types, found immediately outside the cell membrane. It can be tough, flexible, and sometimes rigid. Primarily, it provides the cell with structural support, shape, protection, ...

s and that the

plasma membrane The cell membrane (also known as the plasma membrane or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of a cell from the outside environment (the extr ...

of the gametes is close to the plasma membrane of the cell that surrounds them inside the pollen grain.

Double fertilization in gymnosperms

A far more rudimentary form of double fertilization occurs in the sexual reproduction of an order of gymnosperms commonly known as Gnetales. Specifically, this event has been documented in both ''Ephedra'' and '' Gnetum'', a subset of gnetophytes. In '' Ephedra nevadensis'', a single binucleate sperm cell is deposited into the egg cell. Following the initial fertilization event, the second sperm nucleus is diverted to fertilize an additional egg nucleus found in the egg cytoplasm. In most other seed plants, this second 'ventral canal nucleus' is normally found to be functionally useless. In '' Gnetum gnemon'', numerous free egg nuclei exist in female cytoplasm inside the female

. Succeeding the penetration of the mature female gametophyte by the pollen tube, female cytoplasm and free nuclei move to surround the pollen tube. Released from the binucleate sperm cell are two sperm nuclei which then fuse with free egg nuclei to produce two viable zygotes, a homologous characteristic between families ''Ephedra'' and ''Gnetum''. In both families, the second fertilization event produces an additional diploid

embryo An embryo ( ) is the initial stage of development for 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 sp ...

. This supernumerary embryo is later aborted, leading to the synthesis of only one mature embryo. The additional fertilization product in ''Ephedra'' does not nourish the primary embryo, as the female gametophyte is responsible for nutrient provision. The more primitive process of double fertilization in

gymnosperm The gymnosperms ( ; ) are a group of woody, perennial Seed plant, seed-producing plants, typically lacking the protective outer covering which surrounds the seeds in flowering plants, that include Pinophyta, conifers, cycads, Ginkgo, and gnetoph ...

s results in two diploid nuclei enclosed in the same egg cell. This differs from the

angiosperm Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed within a fruit ...

condition, which results in the separation of the egg cell and

. Comparative molecular research on the genome of ''G. gnemon'' has revealed that gnetophytes are more closely related to

conifers Conifers () are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All e ...

than they are to angiosperms. The rejection of the anthophyte hypothesis, which identifies gnetales and angiosperms are sister taxa, leads to speculation that the process of double fertilization is a product of

convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last comm ...

and arose independently among gnetophytes and angiosperms.

In vitro double fertilization

In vitro double fertilization is often used to study the molecular interactions as well as other aspects of gamete fusion in flowering plants. One of the major obstacles in developing an in vitro double fertilization between male and female gametes is the confinement of the sperm in the pollen tube and the egg in the embryonic sac. A controlled fusion of the egg and sperm has already been achieved with poppy plants. Pollen germination, pollen tube entry, and double fertilization processes have all been observed to proceed normally. In fact, this technique has already been used to obtain seeds in various flowering plants and was named “test-tube fertilization”.

Related structures and functions

Megagametophyte

The female gametophyte, the megagametophyte, that participates in double fertilization in

s which is haploid is called the embryonic sac. This develops within an

, enclosed by the ovary at the base of a

. Surrounding the megagametophyte are (one or) two

integument In biology, an integument is the tissue surrounding an organism's body or an organ within, such as skin, a husk, Exoskeleton, shell, germ or Peel (fruit), rind. Etymology The term is derived from ''integumentum'', which is Latin for "a coverin ...

s, which form an opening called the micropyle. The megagametophyte, which is usually

, originates from the (usually

) megaspore mother cell, also called the megasporocyte. The next sequence of events varies, depending on the particular species, but in most species, the following events occur. The megasporocyte undergoes meiosis, producing four haploid megaspores. Only one of the four resulting megaspores survives. This megaspore undergoes three rounds of mitosis, resulting in seven cells with eight haploid nuclei (the central cell has two nuclei, called the polar nuclei). The lower end of the embryonic sac consists of the haploid egg cell positioned in the middle of two other haploid cells, called synergids. The synergids function in the attraction and guidance of the pollen tube to the megagametophyte through the micropyle. At the upper end of the megagametophyte are three antipodal cells.

Microgametophyte

The male gametophytes, or microgametophytes, that participate in double fertilization are contained within

pollen Pollen is a powdery substance produced by most types of flowers of seed plants for the purpose of sexual reproduction. It consists of pollen grains (highly reduced Gametophyte#Heterospory, microgametophytes), which produce male gametes (sperm ...

grains. They develop within the microsporangia, or pollen sacs, of the anthers on the stamens. Each microsporangium contains diploid

microspore Microspores are land plant spores that develop into male gametophytes, whereas megaspores develop into female gametophytes. The male gametophyte gives rise to sperm cells, which are used for fertilization of an egg cell to form a zygote. Megaspo ...

mother cells, or microsporocytes. Each microsporocyte undergoes meiosis, forming four haploid microspores, each of which can eventually develop into a pollen grain. A microspore undergoes

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 ...

and

cytokinesis Cytokinesis () is the part of the cell division process and part of mitosis during which the cytoplasm of a single eukaryotic cell divides into two daughter cells. Cytoplasmic division begins during or after the late stages of nuclear division ...

in order to produce two separate cells, the generative cell and the tube cell. These two cells in addition to the spore wall make up an immature pollen grain. As the male gametophyte matures, the generative cell passes into the tube cell, and the generative cell undergoes mitosis, producing two sperm cells. Once the pollen grain has matured, the

anthers The stamen (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filamen ...

break open, releasing the pollen. The pollen is carried to the

pistil Gynoecium (; ; : gynoecia) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl (botany), whorl of a flower; it consists ...

of another flower, by wind or animal pollinators, and deposited on the stigma. As the pollen grain germinates, the tube cell produces the pollen tube, which elongates and extends down the long style of the carpel and into the ovary, where its sperm cells are released in the megagametophyte. Double fertilization proceeds from here.

References

{{botany Plant sexuality