Apomixis in flowering plants (angiosperms) [edit]

Agamospermy, asexual reproduction through seeds, occurs in flowering plants through many different [3] mechanisms and a simple hierarchical classification of the different types is not possible. Consequently there are almost as many different usages of terminology for apomixis in angiosperms as there are [4] authors on the subject. For English speakers, Maheshwari 1950 is very influential. German speakers [5] [6] might prefer to consult Rutishauser 1967. Some older text books on the basis of misinformation (that the egg cell in a meiotically unreduced gametophyte can never be fertilized) attempted to reform the terminology to match parthenogenesis as it is used in zoology, and this continues to cause much confusion. Agamospermy occurs mainly in two forms: In gametophytic apomixis, the embryo arises from an unfertilized egg cell (i.e. by parthenogenesis) in a gametophyte that was produced from a cell that did not complete meiosis. In adventitious embryony (sporophytic apomixis), an embryo is formed directly (not from a gametophyte) from nucellus or integument tissue (see nucellar embryony).

Types of apomixis in flowering plants [edit]

Vegetative apomixis in Poa bulbosa;bulbils form instead of flowers

Caribbean Agave producing plantlets on the old flower stem.

Maheshwari

[4]

used the following simple classification of types of apomixis in flowering plants:

Nonrecurrent apomixis: In this type "the megaspore mother cell undergoes the usual meiotic divisions and a haploid embryo sac [megagametophyte] is formed. The new embryo may then arise either from the egg (haploid parthenogenesis) or from some other cell of the gametophyte (haploid apogamy)." The haploid plants have half as many chromosomes as the mother plant, and "the process is not repeated from one generation to another" (which is why it is called nonrecurrent). See also parthenogenesis and apogamybelow. Recurrent apomixis, is now more often called gametophytic apomixis: In this type, the megagametophyte has the same number of chromosomes as the mother plant because meiosis was not completed. It generally arises either from an archesporial cell or from some other part of the nucellus. Adventive embryony, also called sporophytic apomixis, sporophytic budding, or nucellar embryony: Here there may be a megagametophyte in the ovule, but the embryos do not arise from the cells of the gametophyte; they arise from cells of nucellus or the integument. Adventive embryony is important in several species of Citrus, in Garcinia, Euphorbia dulcis, Mangifera indica etc. Vegetative apomixis: In this type "the flowers are replaced by bulbils or other vegetative propagules which frequently germinate while still on the plant". Vegetative apomixis is important in Allium, Fragaria, Agave, and some grasses, among others.
[7]

Types of gametophytic apomixis [edit]

Gametophytic apomixis in flowering plants develops in several different ways. A megagametophyte develops with an egg cell within it that develops into an embryo through parthenogenesis. The central cell of the megagametophyte may require fertilization to form theendosperm, pseudogamous gametophytic apomixis, or in autonomous gametophytic apomixis fertilization is not required. In diplospory (also called generative apospory), the megagametophyte arises from a cell of the archesporium. In apospory (also called somatic apospory), the megagametophyte arises from some other nucellus cell.

Considerable confusion has resulted because diplospory is often defined to involve the megaspore mother cell only, but a number of plant families have a multicellular archesporium and the megagametophyte could originate from another archesporium cell. Diplospory is further subdivided according to how the megagametophyte forms: Allium odorumA. nutans type. The chromosomes double (endomitosis) and then meiosis proceeds in an unusual way, with the chromosome copies pairing up (rather than the original maternal and paternal copies pairing up). Taraxacum type: Meiosis I fails to complete, meiosis II creates two cells, one of which degenerates; three mitotic divisions form the megagametophyte. Ixeris type: Meiosis I fails to complete; three rounds of nuclear division occur without cell-wall formation; wall formation then occurs.

BlumeaElymus types: A mitotic division is followed by degeneration of one cell; three mitotic divisions form the megagametophyte. AntennariaHieracium types: three mitotic divisions form the megagametophyte. EragrostisPanicum types: Two mitotic division give a 4-nucleate megagametophyte, with cell walls to form either three or four cells.

Incidence of apomixis in flowering plants [edit]

Apomixis occurs in at least 33 families of flowering plants, and has evolved multiple times from sexual [8][9] [9] relatives. Apomictic species or individual plants often have a hybrid origin, and are usually polyploid. In plants with both apomictic and meiotic embryology, the proportion of the different types can differ at [7] [7] different times of year, and photoperiod can also change the proportion. It appears unlikely that there are any truly completely apomictic plants, as low rates of sexual reproduction have been found in several [7] species that were previously thought to be entirely apomictic. The genetic control of apomixis can involve a single genetic change that affects all the major developmental components, formation of the megagametophyte, parthenogenesis of the egg cell, and [10] endosperm development. However, the timing of the various developmental processes is critical to successful development of an apomictic seed, and the timing can be affected by multiple genetic [10] factors.

Some terms related to apomixis [edit]

Apomeiosis: "Without meiosis"; usually meaning the production of a meiotically unreduced gametophyte. Parthenogenesis: Development of an embryo directly from an egg cell without fertilization is called parthenogenesis. It is of two types: Haploid parthenogenesis: Parthenogenesis of a normal haploid egg (a meiotically reduced egg) into an embryo is termed haploid parthenogenesis. If the mother plant was diploid, then the haploid embryo that results is monoploid, and the plant that grows from the embryo is sterile. If they are not sterile, they are sometimes useful to plant breeders (especially in potato breeding, see dihaploidy). This type of apomixis has been recorded in Solanum nigram, Lilium spp., Orchis maculate, Nicotiana tabacum etc. Diploid parthenogenesis: When the megagametophyte develops without completing meiosis, so that the megagametophyte and all cells within it are meiotically unreduced (aka diploid, but diploid is an ambiguous term), this is called diploid parthenogenesis, and the plant that develops from the embryo will have the same number of chromosomes as the mother plant. Diploid parthenogenesis is a component process of gametophytic apomixis (see above). Androgenesis and androclinesis are synonyms. These terms are used for two different processes that both have the effect of producing an embryo that has "male inheritance".

The first process is a natural one. It is may also be referred to as male apomixis or paternal apomixis It involves fusion of the male and female gametes and replacement of the female nucleus by the male nucleus. This has been noted as a rare phenomenon in many plants (e.g. Nicotiana and Crepis), and occurs as the regular reproductive method in the Saharan Cypress, Cupressus dupreziana.
[11][12]

The second process that is referred to as androgenesis or androclinesis involves (artificial) culture of haploid plants from anther tissue or microspores.
[13]

Apogamy: Although this term was (before 1908) used for other types of apomixis, and then discarded as too confusing, it is still sometimes used when an embryo develops from a cell of the megagametophyte other than the egg cell. In flowering plants, the cells involved in apogamy would be synergids or antipodal cells. Addition hybrids, called BIII hybrids by Rutishauser: An embryo is formed after a meiotically unreduced egg cell is fertilized. The ploidy level of the embryo is therefore higher than that of the mother plant. This process occurs in some plants that are otherwise apomictic, and may play a significant role in producing tetraploid plants from triploid apomictic mother plants (if they receive pollen from diploids). Because fertilization is involved, this process does not fit the definition of apomixis. Pseudogamy refers to any reproductive process that requires pollination but does not involve male inheritance. It is sometimes used in a restrictive sense to refer to types of apomixis in which the endosperm is fertilized but the embryo is not. A better term for the [13] restrictive sense is centrogamy. Agamospecies, the concept introduced by Gte Turesson: "an apomict population the constituents of which, for morphological, cytological or other reasons, are to be considered [14] as having a common origin," i.e., basically synonymous with "microspecies.
[5]

Parthenocarpy
From Wikipedia, the free encyclopedia

In botany and horticulture, parthenocarpy (literally meaning virgin fruit) is the natural or artificially induced production of fruit without fertilization of ovules. The fruit is thereforeseedless. Stenospermocarpy may also produce apparently seedless fruit, but the seeds are actually aborted while still small. Parthenocarpy (or stenospermocarpy) occasionally occurs as a mutation in nature, but if it affects every flower, then the plant can no longer sexually reproduce but might be able to propagate by vegetative means. However, parthenocarpy of some fruits on a plant may be of value. Up to 20% of the fruits of wild parsnip are parthenocarpic. The seedless wild parsnip fruit are preferred by certain herbivores, and thus serve as a "decoy defense" against seed predation.[1] Utah juniper has a similar defense against bird feeding.[2] Being able to

produce seedless fruit whenpollination is unsuccessful may be an advantage to a plant because it provides food for the plant's seed dispersers. Without a fruit crop, the seed dispersing animals may starve or migrate.

A seedless watermelon.

In some plants, pollination or other stimulation is required for parthenocarpy. This is termed stimulative parthenocarpy. Plants that do not require pollination or other stimulation to produce parthenocarpic fruit have vegetative parthenocarpy. Seedless cucumbers are an example of vegetative parthenocarpy, seedless watermelon is an example of stenospermocarpy. Plants moved from one area of the world to another may not always be accompanied by their pollinating partner and the lack ofpollinators has spurred human cultivation of parthenocarpic varieties. Some parthenocarpic varieties have been developed as genetically modified organisms.
Contents
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1 Commercial importance 2 Misconceptions 3 See also 4 References 5 External links

Commercial importance [edit]

Seedlessness is seen as a desirable trait in edible fruit with hard seeds such as pineapple, banana, orange and grapefruit. Parthenocarpy is also desirable in fruit crops that may be difficult to pollinate or fertilize, such as tomato and summer squash. In dioecious species, such as persimmon, parthenocarpy increases fruit production because staminate trees do not need to be planted to provide pollen. Parthenocarpy is undesirable in nut crops, such as pistachio, where the seed is the edible part. Horticulturists have selected and propagated parthenocarpic cultivars of many plants, including fig, cactus pear

(Opuntia), breadfruit and eggplant. Some plants, such as pineapple, produce seedless fruits when a single cultivar is grown because they are self-infertile. Some cucumbers produce seedless fruit if pollinators are excluded. Strange as it seems, seedless watermelon plants are grown from seeds. The seeds are produced by crossing a diploid parent with a tetraploid parent to produce triploid seeds. When sprayed on flowers, any of the plant hormones gibberellin, auxin and cytokinin can often stimulate the development of parthenocarpic fruit. This is termed artificial parthenocarpy. Plant hormones are seldom used commercially to produce parthenocarpic fruit. Home gardeners sometimes spray their tomatoes with an auxin to assure fruit production. Some parthenocarpic cultivars have been developed as genetically modified organisms.[3] Some parthenocarpic cultivars are of ancient origin. The oldest known cultivated plant is a parthenocarpic fig first grown at least 11,200 years ago.[4] In some climates, normally seeded pear cultivars will produce mainly seedless fruit due to lack of pollination. [5]

Misconceptions [edit]

Most commercial seedless grape cultivars, such as 'Thompson Seedless', are not seedless because of parthenocarpy, but because of stenospermocarpy.[6]

Parthenocarpy is sometimes claimed to be the equivalent of parthenogenesis in animals.[7] That is incorrect because parthenogenesis is a method of asexual reproduction, and parthenocarpy is not, except in rare cases such as pineapple. The plant equivalent of parthenogenesis is apomixis.

See also