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A gametophyte () is one of the two alternating multicellular phases in the life cycles of 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 , 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.
In some bryophyte groups such as many liverworts of the order Marchantiales, the gametes are produced on specialized structures called (or gametangiophores).
In heterospory 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.
In , 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 meiosis 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. have 3 celled pollen grains while Ginkgoales have 4 celled pollen grains. Gnetophyta may have 2 or 3 celled pollen grains depending on the species, and Pinophyta pollen grains vary greatly ranging from single celled to 40 celled. One of these cells is typically a germ cell 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 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 Conifer cone 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 Flagellum 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 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 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 , 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 at that stage, 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. 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). 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 , 2 Synergid, 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.
In heterosporous plants (water ferns, some lycophytes, as well as all gymnosperms and angiosperms), there are two distinct types of sporangium, 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, which becomes the food storage tissue in the seed.
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