Double fertilization or double fertilisation (see spelling differences) is a complex fertilization mechanism of angiosperms. This process involves the fusion of a female gametophyte or megagametophyte, also called the embryonic sac, with two male gametes (sperm). It begins when a pollen grain adheres to the stigmatic surface of the carpel, the female reproductive structure of angiosperm flowers. The pollen grain begins to germinate (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 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, 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 sperm and haploid egg fuse to form a diploid zygote, the process being called syngamy, while the other sperm and the diploid central cell fuse to form a triploid primary endosperm cell (triple fusion). Some plants may form polyploid nuclei. The large cell of the gametophyte will then develop into the endosperm, a nutrient-rich tissue which nourishes the developing embryo. The ovary, surrounding the ovules, develops into the fruit, which protects the seeds and may function to disperse them. [1]
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 , 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 imaging. Some of the genes involved in the migration and fusion process have also been determined. [2]
Evidence of double fertilization in Gnetales, which are non-flowering seed plants, has been reported. [3]
Double fertilization was discovered more than a century ago by Sergei Nawaschin in Kyiv, [4] and Léon Guignard in France. Each made the discovery independently of the other. [5] Lilium martagon and Fritillaria tenella were used in the first observations of double fertilization, which were made using the classical light microscope. 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, 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 walls and that the plasma membrane of the gametes is close to the plasma membrane of the cell that surrounds them inside the pollen grain. [6]
A far more rudimentary form of double fertilization occurs in the sexual reproduction of an order of gymnosperms commonly known as Gnetales. [3] Specifically, this event has been documented in both Ephedra and Gnetum , a subset of gnetophytes. [7] 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. [8] In Gnetum gnemon , numerous free egg nuclei exist in female cytoplasm inside the female gametophyte. 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. [9] In both families, the second fertilization event produces an additional diploid embryo. This supernumerary embryo is later aborted, leading to the synthesis of only one mature embryo. [10] The additional fertilization product in Ephedra does not nourish the primary embryo, as the female gametophyte is responsible for nutrient provision. [9] The more primitive process of double fertilization in gymnosperms results in two diploid nuclei enclosed in the same egg cell. This differs from the angiosperm condition, which results in the separation of the egg cell and endosperm. [11] Comparative molecular research on the genome of G. gnemon has revealed that gnetophytes are more closely related to conifers than they are to angiosperms. [12] [13] [14] 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 and arose independently among gnetophytes and angiosperms. [15]
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. [16] 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”. [17]
The female gametophyte, the megagametophyte, that participates in double fertilization in angiosperms which is haploid is called the embryonic sac. This develops within an ovule, enclosed by the ovary at the base of a carpel. Surrounding the megagametophyte are (one or) two integuments, which form an opening called the micropyle. The megagametophyte, which is usually haploid, originates from the (usually diploid) 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.
The male gametophytes, or microgametophytes, that participate in double fertilization are contained within pollen grains. They develop within the microsporangia, or pollen sacs, of the anthers on the stamens. Each microsporangium contains diploid microspore 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 and cytokinesis 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 break open, releasing the pollen. The pollen is carried to the pistil 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. [18]
A gamete 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. The name gamete was introduced by the German cytologist Eduard Strasburger in 1878.
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 sexual phase in the life cycle of plants and algae. It develops sex organs that produce gametes, haploid sex cells that participate in fertilization to form a diploid zygote which has a double set of chromosomes. Cell division of the zygote results in a new diploid multicellular organism, the second stage in the life cycle known as the sporophyte. The sporophyte can produce haploid spores by meiosis that on germination produce a new generation of gametophytes.
Fertilisation or fertilization, also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a zygote and initiate its development into a new individual organism or offspring. While processes such as insemination or pollination, which happen before the fusion of gametes, are also sometimes informally referred to as fertilisation, these are technically separate processes. The cycle of fertilisation and development of new individuals is called sexual reproduction. During double fertilisation in angiosperms, the haploid male gamete combines with two haploid polar nuclei to form a triploid primary endosperm nucleus by the process of vegetative fertilisation.
In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, fungi and protozoa. They were thought to have appeared as early as the mid-late Ordovician period as an adaptation of early land plants.
Alternation of generations is the predominant type of life cycle in plants and algae. In plants both phases are multicellular: the haploid sexual phase – the gametophyte – alternates with a diploid asexual phase – the sporophyte.
In botany, apomixis is asexual development of seed or embryo without fertilization. However, other definitions include replacement of the seed by a plantlet or replacement of the flower by bulbils.
The egg cell or ovum is the female reproductive cell, or gamete, in most anisogamous organisms. The term is used when the female gamete is not capable of movement (non-motile). If the male gamete (sperm) is capable of movement, the type of sexual reproduction is also classified as oogamous. A nonmotile female gamete formed in the oogonium of some algae, fungi, oomycetes, or bryophytes is an oosphere. When fertilized, the oosphere becomes the oospore.
An archegonium, from the Ancient Greek ἀρχή ("beginning") and γόνος ("offspring"), is a multicellular structure or organ of the gametophyte phase of certain plants, producing and containing the ovum or female gamete. The corresponding male organ is called the antheridium. The archegonium has a long neck canal or venter and a swollen base. Archegonia are typically located on the surface of the plant thallus, although in the hornworts they are embedded.
The gymnosperms are a group of seed-producing plants that includes conifers, cycads, Ginkgo, and gnetophytes, forming the clade Gymnospermae. The term gymnosperm comes from the composite word in Greek: γυμνόσπερμος, literally meaning 'naked seeds'. The name is based on the unenclosed condition of their seeds. The non-encased condition of their seeds contrasts with 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, which are often modified to form cones, or on their own as in yew, Torreya, Ginkgo. The life cycle of a gymnosperm involves alternation of generations, with a dominant diploid sporophyte phase, and a reduced haploid gametophyte phase, which is dependent on the sporophytic phase. The term "gymnosperm" is often used in paleobotany to refer to all non-angiosperm seed plants. In that case, to specify the modern monophyletic group of gymnosperms, the term Acrogymnospermae is sometimes used.
Gametogenesis is a biological process by which diploid or haploid precursor cells undergo cell division and differentiation to form mature haploid gametes. Depending on the biological life cycle of the organism, gametogenesis occurs by meiotic division of diploid gametocytes into various gametes, or by mitosis. For example, plants produce gametes through mitosis in gametophytes. The gametophytes grow from haploid spores after sporic meiosis. The existence of a multicellular, haploid phase in the life cycle between meiosis and gametogenesis is also referred to as alternation of generations.
A sporophyte is the diploid multicellular stage in the life cycle of a plant or alga which produces asexual spores. This stage alternates with a multicellular haploid gametophyte phase.
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, and the female gametophyte in its center. The female gametophyte — specifically termed a megagametophyte — is also called the embryo sac in angiosperms. The megagametophyte produces an egg cell for the purpose of fertilization. The ovule is a small structure present in the ovary. It is attached to the placenta by a stalk called a funicle. The funicle provides nourishment to the ovule. On the basis of the relative position of micropyle, body of the ovule, chalaza and funicle, there are six types of ovules.
The endosperm is a tissue produced inside the seeds of most of the flowering plants following double fertilization. It is triploid in most species, which may be auxin-driven. It surrounds the embryo and provides nutrition in the form of starch, though it can also contain oils and protein. This can make endosperm a source of nutrition in animal diet. For example, wheat endosperm is ground into flour for bread, while barley endosperm is the main source of sugars for beer production. Other examples of endosperm that forms the bulk of the edible portion are coconut "meat" and coconut "water", and corn. Some plants, such as certain orchids, lack endosperm in their seeds.
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. Megaspores are structures that are part of the alternation of generations in many seedless vascular cryptogams, all gymnosperms and all angiosperms. Plants with heterosporous life cycles using microspores and megaspores arose independently in several plant groups during the Devonian period. Microspores are haploid, and are produced from diploid microsporocytes by meiosis.
A gametangium is an organ or cell in which gametes are produced that is found in many multicellular protists, algae, fungi, and the gametophytes of plants. In contrast to gametogenesis in animals, a gametangium is a haploid structure and formation of gametes does not involve meiosis.
Megaspores, also called macrospores, are a type of spore that is present in heterosporous plants. These plants have two spore types, megaspores and microspores. Generally speaking, the megaspore, or large spore, germinates into a female gametophyte, which produces egg cells. These are fertilized by sperm produced by the male gametophyte developing from the microspore. Heterosporous plants include seed plants, water ferns (Salviniales), spikemosses (Selaginellaceae) and quillworts (Isoetaceae).
Plant reproduction is the production of new offspring in plants, which can be accomplished by sexual or asexual reproduction. Sexual reproduction produces offspring by the fusion of gametes, resulting in offspring genetically different from either parent. Asexual reproduction produces new individuals without the fusion of gametes, resulting in clonal plants that are genetically identical to the parent plant and each other, unless mutations occur.
Megagametogenesis is the process of maturation of the female gametophyte, or megagametophyte, in plants. During the process of megagametogenesis, the megaspore, which arises from megasporogenesis, develops into the embryonic sac, which is where the female gamete is housed. These megaspores then develop into the haploid female gametophytes. This occurs within the ovule, which is housed inside the ovary.
A megaspore mother cell, or megasporocyte, is a diploid cell in plants in which meiosis will occur, resulting in the production of four haploid megaspores. At least one of the spores develop into haploid female gametophytes (megagametophytes). The megaspore mother cell arises within the megasporangium tissue.
Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete with a single set of chromosomes combines with another gamete to produce a zygote that develops into an organism composed of cells with two sets of chromosomes (diploid). This is typical in animals, though the number of chromosome sets and how that number changes in sexual reproduction varies, especially among plants, fungi, and other eukaryotes.
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