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Dioecy ( ; ; adj. dioecious, ) is a characteristic of certain that have distinct unisexual individuals, each producing either male or female , either directly (in ) or indirectly (in ). Dioecious reproduction is biparental reproduction. Dioecy has costs, since only the female part of the population directly produces offspring. It is one method for excluding self-fertilization and promoting (outcrossing), and thus tends to reduce the expression of recessive deleterious mutations present in a population. Plants have several other methods of preventing self-fertilization including, for example, , , and self-incompatibility.


In zoology
In zoology, dioecy means that an animal is either male or female, in which case the synonym is more often used. Most species are gonochoric, almost all vertebrate species are gonochoric, and all bird and mammal species are gonochoric. Dioecy may also describe colonies within an , such as the colonies of (Portuguese man-of-war), which may be either dioecious or .


In botany
Land plants () differ from animals in that their life cycle involves alternation of generations. In animals, typically an individual produces of one kind, either or . The gametes have half the number of of the individual producing them, so are . Without further dividing, a sperm and an egg cell fuse to form a that develops into a new individual. In land plants, by contrast, one generation – the generation – consists of individuals that produce haploid rather than haploid . Spores do not fuse, but by dividing repeatedly by to give rise to haploid individuals, the , which produce gametes. A male gamete and a female gamete then fuse to produce a new sporophyte.

In (, and ), the gametophytes are fully independent plants. Seed plant gametophytes are dependent on the sporophyte and develop within the spores, a condition known as endospory. In flowering plants, the male gametophytes develop within grains produced by the sporophyte's , and the female gametophytes develop within produced by the sporophyte's carpels.

The sporophyte generation of a seed plant is called "monoecious" when each sporophyte plant has both kinds of spore-producing organ but in separate flowers or cones. For example, a single of a monoecious species has both functional stamens and carpels, in separate flowers.

The sporophyte generation of seed plants is called when each sporophyte plant has only one kind of spore-producing organ, all of whose spores give rise either to male gametophytes, which produce only male gametes (sperm), or to female gametophytes, which produce only female gametes (egg cells). For example, a single flowering plant sporophyte of a fully dioecious species like has either flowers with functional stamens producing pollen containing male gametes (staminate or 'male' flowers), or flowers with functional carpels producing female gametes (carpellate or 'female' flowers), but not both. There are other, more complex reproductive schemes such as and .

Ilex aquifolium male HC1.JPG|In dioecious holly, some plants only have 'male' flowers with stamens producing pollen. Ilex aquifolium female HC1.JPG|Other holly plants only have 'female' flowers that produce ovules. Tulip Tulipa clusiana 'Lady Jane' Rock Ledge Flower 2000px.jpg|Each bisexual (perfect) tulip flower has both stamens and carpels.

Slightly different terms, dioicous and , may be used for the generation of non-vascular plants, although dioecious and monoecious are also used. A dioicous gametophyte either produces only male gametes (sperm) or produces only female gametes (egg cells). About 60% of liverworts are dioicous.

(2025). 9780521700733, Cambridge University Press.

Dioecy occurs in a wide variety of plant groups. Examples of dioecious plant species include , , and . As its specific name implies, the perennial stinging nettle is dioecious,

(2025). 9781527226302, C & M Floristics.
while the annual nettle is monoecious. Dioecious are predominant in environments.
(2020). 9789811542107, Springer Nature. .

About 65% of species are dioecious, but almost all conifers are monoecious. In gymnosperms, the sexual systems dioecy and monoecy are strongly correlated with the mode of pollen dispersal, monoecious species are predominantly wind dispersed () and dioecious species animal-dispersed ().

About 6 percent of species are entirely dioecious and about 7% of angiosperm contain some dioecious species. Dioecy is more common in , and species. In most dioecious plants, whether male or female gametophytes are produced is determined genetically, but in some cases it can be determined by the environment, as in species.

(2019). 9781108499859, Cambridge University Press. .

Certain , such as some species of , are dioecious.Maggs, C.A. and Hommersand, M.H. 1993. Seaweeds of the British Isles Volume 1 Rhodophyta Part 3A Ceramiales. The Natural History Museum, London. Dioecy is prevalent in the brown algae () and may have been the ancestral state in that group.


Evolution of dioecy
In plants, dioecy has evolved independently multiple times either from hermaphroditic species or from monoecious species. A previously untested hypothesis is that this reduces inbreeding; dioecy has been shown to be associated with increased genetic diversity and greater protection against deleterious mutations. Regardless of the evolutionary pathway the intermediate states need to have fitness advantages compared to cosexual flowers in order to survive.
(2018). 9780190882686, Oxford University Press. .

Dioecy evolves due to male or female sterility,

(1999). 9780732944391, Macmillan Education AU. .
although it is unlikely that mutations for male and female sterility occurred at the same time.
(2015). 9783319212548, Springer. .
In angiosperms unisexual flowers evolve from bisexual ones.
(2020). 9783030460129, Springer Nature. .
Dioecy occurs in almost half of plant families, but only in a minority of genera, suggesting recent evolution.
(2014). 9781134263509, Routledge. .
For 160 families that have dioecious species, dioecy is thought to have evolved more than 100 times.

In the family , dioecy is likely the ancestral sexual system.

(2020). 9781789241907, CABI. .


From monoecy
Dioecious flowering plants can evolve from monoecious ancestors that have flowers containing both functional stamens and functional carpels. Some authors argue monoecy and dioecy are related.
(2019). 9781439844366, CRC Press. .

In the genus , since there is a distribution of sexual systems, it has been postulated that dioecy evolved from monoecy

(2000). 9780643099296, Csiro Publishing. .
through mainly from mutations that resulted in male sterility.
(2016). 9780128004265, Academic Press. .
However, since the ancestral state is unclear, more work is needed to clarify the evolution of dioecy via monoecy.


From hermaphroditism
Dioecy usually evolves from hermaphroditism through but may also evolve through , through
(2019). 9783319941394, Springer. .
or through . In the , dioecy may have evolved independently from hermaphroditism at least 5 or 9 times. The reverse transition, from dioecy back to hermaphroditism has also been observed, both in Asteraceae and in bryophytes, with a frequency about half of that for the forward transition.
(2013). 9789400773479, Springer Science & Business Media. .

In , since there is no monoecy, it is suggested that dioecy evolved through gynodioecy.


In mycology
Very few dioecious fungi have been discovered.
9788176487375, APH Publishing. .

Monoecy and dioecy in refer to the donor and recipient roles in mating, where a nucleus is transferred from one haploid hypha to another, and the two nuclei then present in the same cell merge by to form a . The definition avoids reference to male and female reproductive structures, which are rare in fungi. An individual of a dioecious fungal species not only requires a partner for mating, but performs only one of the roles in nuclear transfer, as either the donor or the recipient. A monoecious fungal species can perform both roles, but may not be self-compatible.


Adaptive benefit
Dioecy has the demographic disadvantage compared with hermaphroditism that only about half of reproductive adults are able to produce offspring. Dioecious species must therefore have fitness advantages to compensate for this cost through increased survival, growth, or reproduction. Dioecy excludes self-fertilization and promotes (outcrossing), and thus tends to reduce the expression of recessive deleterious mutations present in a population. In trees, compensation is realized mainly through increased seed production by females. This in turn is facilitated by a lower contribution of reproduction to population growth, which results in no demonstrable net costs of having males in the population compared to being hermaphroditic. Dioecy may also accelerate or retard lineage diversification in . Dioecious lineages are more diversified in certain genera, but less in others. An analysis suggested that dioecy neither consistently places a strong brake on diversification, nor strongly drives it.


See also


Bibliography

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