Gametophyte

Several gametophytes growing in a terrarium.

Pine gametophyte (outside) surrounding the embryo (inside)

A gametophyte is a stage in the life cycle of plants and algae that undergo alternation of generations. 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 in which each cell has two sets 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 function of which is to produce haploid spores by meiosis.

Algae

In some multicellular green algae (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.^[1]

Land plants

In land plants, anisogamy is universal. As in animals, female and male gametes are called, respectively, eggs and sperm. Either the sporophyte or the gametophyte may be reduced (heteromorphic).

Bryophytes

In bryophytes (mosses, liverworts, 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 typically attached to the gametophytes and dependent on them.^[2] When a moss spore germinates it grows to produce a filament of cells (called the protonema). The mature gametophyte of mosses develops into leafy shotos that produce sex organs (gametangia) that produce gametes. Eggs develop in archegonia and sperm in antheridia.^[3]

In some bryophyte groups such as many liverworts of the order Marchantiales, the gametes are produced on specialized structures called gametophores (or gametangiophores).

Ferns

In most ferns, gametophytes are photosynthetic free living organism called a prothallus. However in some groups, notably the clade that includes Ophioglossaceae and Psilotaceae, gametophytes are subterranean and subsist by forming mycotrophic relationships with fungi. In leptosporangiate ferns such as Dryopteris, the gametophyte is a free-living, autotrophic prothallus that maintains the sporophyte during its early multicellular development.

Lycophytes

Extant lycophytes produce several different types of gametophytes. In the families Lycopodiaceae and Huperziaceae, gametophytes are subterranean and mycotrophic, deriving nutrients from symbiosis with fungi. Isoetes and Selaginella, which are heterosporous, megagametophytes develop inside the megaspores, which crack 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 giant arborescent clubmosses, Lepidodendron and Lepidostrobus.^[4]

Seed plants

By contrast, in seed plants (gymnosperms and angiosperms), gametophytes develop into multicellular organisms while still enclosed within the sporangium.^[5]

Vascular plants 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 are normally bisexual, capable of producing both sperm and eggs. In heterosporous vascular plants (plants that produce both microspores and megaspores), the gametophyte develops endosporically, within the spore wall. These gametophytes are unisexual, producing either sperm or eggs but not both.

All vascular plants are sporophyte dominant, and a trend toward smaller and more sporophyte-dependent female gametophytes is evident as land plants evolved towards reproduction by seeds.^[6]

Heteromorphy

In plants with heteromorphic gametophytes, 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. Gametophytes which produce egg and sperm on separate plants are termed Dioicous.

In heterosporous plants (water ferns, some lycophytes, as well as all gymnosperms and angiosperms), there are two distinct sporangia, each of which produces a single kind of spore and 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 the seed plants, the microgametophyte is called pollen. Seed plant microgametophytes consists of two or three 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 grain) 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.^[7]

In angiosperms, the megagametophyte is reduced to only a few nuclei and 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 endosperm, which becomes the food storage tissue in the seed.

See also

• Sporophyte
• Alternation of generations
• Archegonium
• Antheridium

References

1. Sadava, David; Hillis, David; Heller, H. Craig; Berenbaum, May (2012). Life: The Science of Biology, Volume 1 (10th ed.). Macmillan. ISBN 978-1464141225.
2. Budke, J.M., Goffinet, B. and Jones, C.S. (2013):Dehydration protection provided by a maternal cuticle improves offspring fitness in the moss Funaria hygrometrica. Annals of Botany doi:10.1093/aob/mct033
3. Ralf Reski (1998): Development, genetics and molecular biology of mosses. In: Botanica Acta 111, pp 1-15.
4. Brack-Hanes, S.D. (1978). "On the megagametophytes of two Lepidodendracean cones.". Botanical Gazette 139: 140–146.
5. C.Michael Hogan (2010): Fern. Encyclopedia of Earth. National council for Science and the Environment. Washington, DC
6. Stewart, W.N.; Rothwell, G.W. Palaeobotany and the evolution of plants, second edition. Cambridge, U.K.: Cambridge University press. ISBN 0521382947.
7. "Vascular Plants :: Description". Digimuse.nmns.edu.tw. Retrieved 2014-07-13.