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A gametophyte () is one of the two alternating multicellular phases in the life cycles of
plant Plants are predominantly photosynthetic eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota o ...
s and
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organism In biology Biology is the scientific study of life. It is a natural science with a broad scope but has seve ...
. It is a haploid multicellular organism that develops from a haploid
spore In biology Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells tha ...
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
gamete A gamete (; , ultimately ) is a haploid cell that fuses with another haploid cell during fertilization in organism In biology Biology is the scientific study of life. It is a natural science with a broad scope but has s ...
s, haploid sex cells that participate in fertilization to form a
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible allele An allele (, ; ; modern formation from Greek ἄλλος ''állos'', "other") is a variation of the same sequence of nucleotide ...
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 A sporophyte () is the diploid multicellular stage in the life cycle of a plant Plants are predominantly photosynthetic eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many un ...
. The sporophyte can produce haploid spores by
meiosis Meiosis (; , since it is a reductional division) is a special type of cell division Cell division is the process by which a parent cell divides into two daughter cells. Cell division usually occurs as part of a larger cell cycle T ...
that on germination produce a new generation of gametophytes.


Algae

In some multicellular
green algae The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/ Streptophyta. The land plants ( Embryophytes) have emerged deep in the Charophyte alg ...
('' Ulva lactuca'' is one example), red algae and brown algae, sporophytes and gametophytes may be externally indistinguishable (isomorphic). In '' Ulva'' the gametes are isogamous, all of one size, shape and general morphology.


Land plants

In land plants,
anisogamy Different forms of anisogamy: A) anisogamy of motile cells, B) egg_cell.html"_;"title="oogamy_(egg_cell">oogamy_(egg_cell_and_sperm_cell),_C)_anisogamy_of_non-motile_cells_(egg_cell_and_spermatia)..html" ;"title="egg_cell_and_sperm_cell.html" ;" ...
is universal. As in animals, female and male gametes are called, respectively, ''eggs'' and ''sperm.'' In extant land plants, either the sporophyte or the gametophyte may be reduced (heteromorphic). No extant gametophytes have
stoma In botany, a stoma (from Greek ''στόμα'', "mouth", plural "stomata"), also called a stomate (plural "stomates"), is a pore found in the epidermis of leaves, stems, and other organs, that controls the rate of gas exchange. The pore is ...
ta, but they have been found on fossil species like the early Devonian '' Aglaophyton'' from the Rhynie chert. Other fossil gametophytes found in the Rhynie chert shows they were much more developed than present forms, resembling the sporophyte in having a well-developed conducting strand, a cortex, an epidermis and a cuticle with stomata, but were much smaller.


Bryophytes

In bryophytes (
mosses Mosses are small, non-vascular flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitat ...
,
liverworts The Marchantiophyta () are a division of non-vascular land plant The Embryophyta (), or land plants, are the most familiar group of green plant Plants are predominantly photosynthetic eukaryote Eukaryotes () are organisms wh ...
, and hornworts), the gametophyte is the most visible stage of the life cycle. The bryophyte gametophyte is longer lived, nutritionally independent, and the sporophytes are attached to the gametophytes and dependent on them. When a moss spore germinates it grows to produce a filament of cells (called the protonema). The mature gametophyte of mosses develops into leafy shoots that produce sex organs ( gametangia) that produce gametes. Eggs develop in archegonia and sperm in antheridia. In some bryophyte groups such as many liverworts of the order Marchantiales, the gametes are produced on specialized structures called gametophores (or gametangiophores).


Vascular plants

All vascular plants are sporophyte dominant, and a trend toward smaller and more sporophyte-dependent female gametophytes is evident as land plants evolved reproduction by seeds. Those vascular plants, such as clubmosses and many ferns, that produce only one type of spore are said to be homosporous. They have exosporic gametophytes — that is, the gametophyte is free-living and develops outside of the spore wall. Exosporic gametophytes can either be bisexual, capable of producing both sperm and eggs in the same thallus ( monoicous), or specialized into separate male and female organisms (dioicous). In heterosporous vascular plants (plants that produce both microspores and megaspores), the gametophytes develop endosporically (within the spore wall). These gametophytes are dioicous, producing either sperm or eggs but not both.


Ferns

In most
fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spore In biology Biology is the scientific study of life. It is a natural science wit ...
s, for example, in the
leptosporangiate fern The Polypodiidae, commonly called leptosporangiate ferns, formerly Leptosporangiatae, are one of four subclasses of fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloe ...
'' Dryopteris'', the gametophyte is a photosynthetic free living
autotroph An autotroph or primary producer is an organism that produces complex organic compound In chemistry, organic compounds are generally any chemical compounds that contain carbon- hydrogen or carbon-carbon bonds. Due to carbon's ability to ...
ic organism called a prothallus that produces gametes and maintains the sporophyte during its early multicellular development. However, in some groups, notably the clade that includes Ophioglossaceae and
Psilotaceae Psilotaceae is a family of fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spore In biology Biology is the scientific study of ...
, the gametophytes are subterranean and subsist by forming mycotrophic relationships with fungi. Homosporous ferns secrete a chemical called antheridiogen.


Lycophytes

Extant lycophytes produce two different types of gametophytes. In the homosporous families Lycopodiaceae and Huperziaceae, spores germinate into bisexual free-living, subterranean and mycotrophic gametophytes that derive nutrients from symbiosis with fungi. In '' Isoetes'' and '' Selaginella'', which are heterosporous, microspores and megaspores are dispersed from sporangia either passively or by active ejection. Microspores produce microgametophytes which produce sperm. Megaspores produce reduced megagametophytes inside the spore wall. At maturity, the megaspore cracks open at the trilete suture to allow the male gametes to access the egg cells in the archegonia inside. The gametophytes of ''Isoetes'' appear to be similar in this respect to those of the extinct
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonif ...
arborescent lycophytes ''Lepidodendron'' and ''Lepidostrobus''.


Seed plants

The seed plant gametophyte life cycle is even more reduced than in basal taxa (ferns and lycophytes). Seed plant gametophytes are not independent organisms and depend upon the dominant sporophyte tissue for nutrients and water. With the exception of mature
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gameto ...
, if the gametophyte tissue is separated from the sporophyte tissue it will not survive. Due to this complex relationship and the small size of the gametophyte tissue—in some situations single celled—differentiating with the human eye or even a microscope between seed plant gametophyte tissue and sporophyte tissue can be a challenge. While seed plant gametophyte tissue is typically composed of mononucleate haploid cells (1 x n), specific circumstances can occur in which the ploidy does vary widely despite still being considered part of the gametophyte. In gymnosperms, the male gametophytes are produced inside microspores within the microsporangia located inside male cones or microstrobili. In each microspore, a single gametophyte is produced, consisting of four haploid cells produced by meiotic division of a diploid microspore mother cell.  At maturity, each microspore-derived gametophyte becomes a pollen grain. During its development, the water and nutrients that the male gametophyte requires are provided by the sporophyte tissue until they are released for pollination. The cell number of each mature pollen grain varies between the gymnosperm orders.
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 ma ...
ophyta have 3 celled pollen grains while Ginkgophyta have 4 celled pollen grains. Gnetophyta may have 2 or 3 celled pollen grains depending on the species, and Coniferophyta pollen grains vary greatly ranging from single celled to 40 celled. One of these cells is typically a
germ cell Germ or germs may refer to: Science * Germ (microorganism), an informal word for a pathogen * Germ cell, cell that gives rise to the gametes of an organism that reproduces sexually * Germ layer, a primary layer of cells that forms during emb ...
and other cells may consist of a single tube cell which grows to form the pollen tube, sterile cells, and/or prothallial cells which are both vegetative cells without an essential reproductive function. After
pollination Pollination is the transfer of pollen from an anther The stamen ( plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower A flower, sometimes known as a bloom or blossom, is the reproductive struc ...
is successful, the male gametophyte continues to develop. If a tube cell was not developed in the microstrobilus, one is created after pollination via mitosis. The tube cell grows into the diploid tissue of the female cone and may branch out into the megastrobilus tissue or grow straight towards the egg cell. The megastrobilus sporophytic tissue provides nutrients for the male gametophyte at this stage. In some gymnosperms, the tube cell will create a direct channel from the site of pollination to the egg cell, in other gymnosperms, the tube cell will rupture in the middle of the megastrobilus sporophyte tissue. This occurs because in some gymnosperm orders, the germ cell is nonmobile and a direct pathway is needed, however, in Cycadophyta and Ginkgophyta, the germ cell is mobile due to flagella being present and a direct tube cell path from the pollination site to the egg is not needed. In most species the germ cell can be more specifically described as a sperm cell which mates with the egg cell during fertilization, though that is not always the case. In some Gnetophyta species, the germ cell will release two sperm nuclei that undergo a rare gymnosperm double fertilization process occurring solely with sperm nuclei and not with the fusion of developed cells. After fertilization is complete in all orders, the remaining male gametophyte tissue will deteriorate. The female gametophyte in gymnosperms differs from the male gametophyte as it spends its whole life cycle in one organ, 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 ...
located inside the megastrobilus or female cone. Similar to the male gametophyte, the female gametophyte normally is fully dependent on the surrounding sporophytic tissue for nutrients and the two organisms cannot be separated. However, the female gametophytes of ''
Ginkgo biloba ''Ginkgo biloba'', commonly known as ginkgo or gingko ( ), also known as the maidenhair tree, is a species of tree native to China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's m ...
'' do contain chlorophyll and can produce some of their own energy, though, not enough to support itself without being supplemented by the sporophyte. The female gametophyte forms from a diploid megaspore that undergoes meiosis and starts being singled celled. The size of the mature female gametophyte varies drastically between gymnosperm orders. In Cycadophyta, Ginkgophyta, Coniferophyta, and some Gnetophyta, the single celled female gametophyte undergoes many cycles of mitosis ending up consisting of thousands of cells once mature. At a minimum, two of these cells are egg cells and the rest are haploid
somatic cell A somatic cell (from Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world Ancient history is a time period from the beginning of writing and recorded human history ...
s, but more egg cells may be present and their ploidy, though typically haploid, may vary. In select Gnetophyta, the female gametophyte stays singled celled. Mitosis does occur, but no cell divisions are ever made. This results in the mature female gametophyte in some Gnetophyta having many free nuclei in one cell. Once mature, this single celled gametophyte is 90% smaller than the female gametophytes in other gymnosperm orders. After fertilization, the remaining female gametophyte tissue in gymnosperms serves as the nutrient source for the developing zygote (even in Gnetophyta where the diploid zygote cell is much smaller then, and for a while lives within the single celled gametophyte). The precursor to the male angiosperm gametophyte is a diploid microspore mother cell located inside the
anther The stamen ( plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division An ...
. Once the microspore undergoes meiosis, 4 haploid cells are formed, each of which is a singled celled male gametophyte. The male gametophyte will develop via one or two rounds of mitosis inside the anther. This creates a 2 or 3 celled male gametophyte which becomes known as the pollen grain once dehiscing occurs. One cell is the tube cell, and the remaining cell/cells are the sperm cells. The development of the three celled male gametophyte prior to dehiscing has evolved multiple times and is present in about a third of angiosperm species allowing for faster fertilization after pollination. Once pollination occurs, the tube cell grows in size and if the male gametophyte is only 2 cells at this stage, the single sperm cell undergoes mitosis to create a second sperm cell. Just like in gymnosperms, the tube cell in angiosperms obtains nutrients from the sporophytic tissue, and may branch out into the pistil tissue or grow directly towards the ovule. Once double fertilization is completed, the tube cell and other vegetative cells, if present, are all that remains of the male gametophyte and soon degrade. The female gametophyte of angiosperms develops in the ovule (located inside the female or hermaphrodite
flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechan ...
). Its precursor is a diploid megaspore that undergoes meiosis which produces four haploid daughter cells. Three of these independent gametophyte cells degenerate and the one that remains is the gametophyte mother cell which normally contains one nucleus. In general, it will then divide by mitosis until it consists of 8 nuclei separated into 1 egg cell, 3 antipodal cells, 2 synergid cells, and a central cell that contains two nuclei. In select angiosperms, special cases occur in which the female gametophyte is not 7 celled with 8 nuclei. On the small end of the spectrum, some species have mature female gametophytes with only 4 cells, each with one nuclei. Conversely, some species have 10-celled mature female gametophytes consisting of 16 total nuclei. Once double fertilization occurs, the egg cell becomes the zygote which is then considered sporophyte tissue. Scholars still disagree on whether the fertilized central cell is considered gametophyte tissue. Some botanists consider this endospore as gametophyte tissue with typically 2/3 being female and 1/3 being male, but as the central cell before double fertilization can range from 1n to 8n in special cases, the fertilized central cells range from 2n (50% male/female) to 9n (1/9 male, 8/9th female). However, other botanists consider the fertilized endospore as sporophyte tissue. Some believe it is neither.


Heterospory

In heterosporic plants, there are two distinct kinds of gametophytes. Because the two gametophytes differ in form and function, they are termed ''heteromorphic'', from ''hetero''- "different" and ''morph'' "form". The egg-producing gametophyte is known as a megagametophyte, because it is typically larger, and the sperm producing gametophyte is known as a microgametophyte. Species which produce egg and sperm on separate gametophytes plants are termed dioicous, while those that produce both eggs and sperm on the same gametophyte are termed monoicous. In heterosporous plants (water ferns, some lycophytes, as well as all gymnosperms and angiosperms), there are two distinct types of sporangia, each of which produces a single kind of spore that germinates to produce a single kind of gametophyte. However, not all heteromorphic gametophytes come from heterosporous plants. That is, some plants have distinct egg-producing and sperm-producing gametophytes, but these gametophytes develop from the same kind of spore inside the same sporangium; '' Sphaerocarpos'' is an example of such a plant. In seed plants, the microgametophyte is called pollen. Seed plant microgametophytes consists of several (typically two to five) cells when the pollen grains exit the sporangium. The megagametophyte develops within the megaspore of extant seedless vascular plants and within the megasporangium in a cone or flower in seed plants. In seed plants, the microgametophyte (pollen) travels to the vicinity of the egg cell (carried by a physical or animal vector) and produces two sperm by mitosis. In gymnosperms, the megagametophyte consists of several thousand cells and produces one to several archegonia, each with a single egg cell. The gametophyte becomes a food storage tissue in the seed. In angiosperms, the megagametophyte is reduced to only a few cells, and is sometimes called the embryo sac. A typical embryo sac contains seven cells and eight nuclei, one of which is the egg cell. Two nuclei fuse with a sperm nucleus to form the primary endospermic nucleus which develops to form triploid
endosperm The endosperm is a tissue produced inside the seed A seed is an embryonic plant Plants are predominantly photosynthetic eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many ...
, which becomes the food storage tissue in the seed.


See also

* * * *


References


Further reading

* Roig-Villanova, Irma; Bou, Jordi; Sorin, Céline; Devlin, Paul F.; Martínez-García, Jaime F.
Identification of Primary Target Genes of Phytochrome Signaling. Early Transcriptional Control during Shade Avoidance Responses in Arabidopsis
. ''Plant Physiology'', 141, 1, 2006-05, pàg. 85–96. DOI: 10.1104/pp.105.076331. ISSN: 0032–0889. * Cucinotta, Mara; Colombo, Lucia; Roig-Villanova, Irma (2014).
Ovule development, a new model for lateral organ formation
. ''Frontiers in Plant Science''. 5. doi:10.3389/fpls.2014.00117. ISSN 1664-462X. PMC 3973900.
PMID PubMed is a free search engine A search engine is a software system designed to carry out web searches. They search the World Wide Web in a systematic way for particular information specified in a textual web search query. The searc ...
 24723934. {{Botany Plant morphology Plant anatomy Plant reproduction