View source for Biology:Dioecy

{{Short description|Having distinct male and female organisms}}

'''Dioecy''' ({{IPAc-en|d|aɪ|ˈ|iː|s|i}} {{respell|dy|EE|see}};<ref>{{Cite dictionary |url=http://www.lexico.com/definition/dioecy |archive-url=https://web.archive.org/web/20211221101905/https://www.lexico.com/definition/dioecy |url-status=dead |archive-date=December 21, 2021 |title=dioecy |dictionary=Lexico UK English Dictionary |publisher=[[Organization:Oxford University Press|Oxford University Press]]}}</ref> {{etymology|grc|''{{lang|grc|{{wikt-lang|grc|διοικία}}}}'' {{grc-transl|διοικία}}|two households}}; adj. '''dioecious''', {{IPAc-en|d|aɪ|ˈ|iː|ʃ|(|i|)|ə|s}} {{respell|dy|EE|sh(ee|)əs}}<ref>{{Cite dictionary |url=http://www.lexico.com/definition/dioecious |archive-url=https://web.archive.org/web/20210412183330/https://www.lexico.com/definition/dioecious |url-status=dead |archive-date=April 12, 2021 |title=dioecious |dictionary=Lexico UK English Dictionary UK English Dictionary |publisher=[[Organization:Oxford University Press|Oxford University Press]]}}</ref><ref>{{cite web |title=diœcious, ''adj''. |work=Oxford English Dictionary online |publisher=Oxford University Press |url=https://www.oed.com/view/Entry/53093 |access-date=2021-12-21}}</ref>) is a characteristic of certain species that have distinct unisexual individuals, each producing either male or female [[Biology:Gamete|gamete]]s, either directly (in animals) or indirectly (in [[Biology:Spermatophyte|seed plants]]). '''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 [[Biology:Allogamy|allogamy]] (outcrossing), and thus tends to reduce the expression of recessive deleterious mutations present in a population. [[Biology:Plant|Plant]]s have several other methods of preventing self-fertilization including, for example, dichogamy, [[Biology:Herkogamy|herkogamy]], and self-incompatibility.

==In zoology==
[[File:Physalia.jpg|thumb|right|''Physalia physalis'', [[Biology:Portuguese man o' war|Portuguese man o' war]], is a dioecious colonial marine animal; the reproductive medusae within the colony are all of the same sex.<ref>{{cite web |url=http://animaldiversity.ummz.umich.edu/accounts/Physalia_physalis/ |title=Animal Diversity Web |access-date=27 April 2014}}</ref>]]
{{Further|Gonochorism}}<!-- The term dioecy is usually applied to plants while gonochoric is usually applied to animals. If you plan on adding more information regarding dioecy and animals go edit on [[Biology:Gonochorism|gonochorism]]. -->
In zoology, dioecy means that an animal is either male or female, in which case the synonym gonochory is more often used.<ref name="Kliman-2016" />{{Page needed|date=August 2021}} For example, most [[Biology:Animal|animal]] species are gonochoric, almost all vertebrate species are gonochoric, and all bird and mammal species are gonochoric.<ref>{{cite journal|author=David, J.R.|year=2001|title=Evolution and development: some insights from evolutionary theory|journal=Anais da Academia Brasileira de Ciências|volume=73|issue=3|pages=385–395|doi=10.1590/s0001-37652001000300008|pmid=11600899|doi-access=free}}</ref> Dioecy may also describe colonies within a species, such as the colonies of [[Biology:Siphonophorae|Siphonophorae]] (Portuguese man-of-war), which may be either dioecious or [[Biology:Monoecy|monoecious]].<ref>{{cite journal |author1=Dunn, C.W. |author2=Pugh, P.R. |author3=Haddock, S.H.D. |year=2005 |title=Molecular Phylogenetics of the Siphonophora (Cnidaria), with Implications for the Evolution of Functional Specialization |journal=Systematic Biology |volume=54 |issue=6 |pages=916–935 |doi=10.1080/10635150500354837 |pmid=16338764|doi-access= }}</ref>

==In botany==
Land plants ([[Biology:Embryophyte|embryophyte]]s) differ from animals in that their [[Biology:Biological life cycle|life cycle]] involves [[Biology:Alternation of generations|alternation of generations]]. In animals, typically an individual produces [[Biology:Gamete|gamete]]s of one kind, either [[Biology:Sperm|sperm]] or [[Biology:Egg cell|egg cell]]s. The gametes have half the number of [[Biology:Chromosome|chromosome]]s of the individual producing them, so are [[Biology:Haploid|haploid]]. Without further dividing, a sperm and an egg cell fuse to form a [[Biology:Zygote|zygote]] that develops into a new individual. In land plants, by contrast, one generation – the [[Biology:Sporophyte|sporophyte]] generation – consists of individuals that produce haploid [[Biology:Spore|spore]]s rather than haploid [[Biology:Gamete|gamete]]s. Spores do not fuse, but [[Biology:Germination|germinate]] by dividing repeatedly by [[Biology:Mitosis|mitosis]] to give rise to haploid multicellular individuals, the [[Biology:Gametophyte|gametophyte]]s, which produce gametes. A male gamete and a female gamete then fuse to produce a new diploid sporophyte.{{sfnp|Mauseth|2014|pp=204–205}}

[[File:Alternation of generations simpler still.svg|thumb|center|upright=1.25|Alternation of generations in plants: the sporophyte generation produces spores that give rise to the gametophyte generation, which produces gametes that fuse to give rise to a new sporophyte generation.]]

In bryophytes ([[Biology:Moss|moss]]es, [[Biology:Marchantiophyta|liverworts]] and [[Biology:Hornwort|hornwort]]s), the gametophytes are fully independent plants.{{sfnp|Mauseth|2014|p=487}} Seed plant gametophytes are dependent on the sporophyte and develop within the spores, a condition known as [[Biology:Endospory in plants|endospory]]. In flowering plants, the male gametophytes develop within [[Biology:Pollen|pollen]] grains produced by the sporophyte's [[Biology:Stamen|stamen]]s, and the female gametophytes develop within [[Biology:Ovule|ovule]]s produced by the sporophyte's [[Biology:Gynoecium#Carpels|carpels]].{{sfnp|Mauseth|2014|pp=204–205}}

The sporophyte generation of a seed plant is called "[[Biology:Glossary of botanical terms#monoecious|monoecious]]" when each sporophyte plant has both kinds of spore-producing organ but in separate flowers or cones. For example, a single [[Biology:Flowering plant|flowering plant]] of a monoecious species has both functional stamens and carpels, in separate flowers.{{sfnp|Mauseth|2014|p=218}}

The sporophyte generation of seed plants is called "[[Biology:Glossary of botanical terms#dioecious|dioecious]]" 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 [[Biology:Ilex aquifolium|holly]] 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.{{sfnp|Mauseth|2014|p=218}}<ref name=Hick01>{{Cite book |last1=Hickey |first1=M. |last2=King |first2=C. |year=2001 |title=The Cambridge Illustrated Glossary of Botanical Terms |publisher=Cambridge University Press |name-list-style=amp}}</ref>  (See [[Biology:Plant reproductive morphology#Flowering plants|Plant reproductive morphology]] for further details, including more complex cases, such as [[Biology:Gynodioecy|gynodioecy]] and [[Biology:Androdioecy|androdioecy]].)
<gallery mode=packed heights=160px>
Ilex aquifolium male HC1.JPG|In dioecious holly, some plants only have 'male' flowers with functional stamens that produce 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 monoecious tulip flower has both pollen-producing stamens and carpels containing ovules.
</gallery>

Slightly different terms, ''dioicous'' and ''[[Biology:Monoicous|monoicous]]'', may be used for the [[Biology:Gametophyte|gametophyte]] generation, although ''dioecious'' and ''monoecious'' are also used.<ref name=Lepp07>{{Cite web |last1=Lepp |first1=Heino |date=2007 |title=Case studies : -oicy : Dioicous, dioecious, monoicous and monoecious |website=Australian Bryophytes |publisher=Australian National Botanic Gardens and Australian National Herbarium |url=https://www.anbg.gov.au/bryophyte/case-studies/-oicy.html |access-date=2021-06-21 }}</ref><ref>{{cite book|author=Stearn, W.T.|year=1992|title=Botanical Latin: History, grammar, syntax, terminology and vocabulary, Fourth edition|publisher=David and Charles}}</ref> A dioicous gametophyte either produces only male gametes (sperm) or produces only female gametes (egg cells). About 60% of liverworts are dioicous.<ref name=Vanderpoorten>{{cite book |vauthors=Vanderpoorten A,Goffinet B |date=2009 |title=Introduction to bryophytes |chapter=Liverworts |publisher=Cambridge University Press |location=Cambridge, UK |isbn=978-0-521-70073-3 }}</ref>{{rp|52}}

Dioecy occurs in a wide variety of plant groups. Examples of dioecious plant species include [[Biology:Ginkgo|ginkgo]]s, [[Biology:Willow|willow]]s, [[Biology:Cannabis|cannabis]] and [[Biology:Milicia excelsa|African teak]]. As its specific name implies, the perennial stinging nettle ''[[Biology:Urtica dioica|Urtica dioica]]'' is dioecious,<ref name=Stace>{{cite book|last=Stace|first=C. A.|year=2019|title=New Flora of the British Isles|edition=Fourth|publisher=C & M Floristics|location = Middlewood Green, Suffolk, U.K.| isbn=978-1-5272-2630-2}}</ref>{{rp|305}} while the annual nettle ''[[Biology:Urtica urens|Urtica urens]]'' is monoecious.<ref name=Stace/>{{rp|305}} Dioecious [[Earth:Flora|flora]] are predominant in tropical environments.<ref>{{Cite book|last1=Tandon|first1=Rajesh|url=https://books.google.com/books?id=WED2DwAAQBAJ&q=dioecy+tropical+climates&pg=PA179|title=Reproductive Ecology of Flowering Plants: Patterns and Processes|last2=Shivanna|first2=K. R.|last3=Koul|first3=Monika|date=2020-08-07|publisher=Springer Nature|isbn=978-981-15-4210-7|pages=179|language=en}}</ref>

About 65% of [[Biology:Gymnosperm|gymnosperm]] species are dioecious,<ref>{{Cite journal|date=2018-09-01|title=Sexual systems in gymnosperms: A review|url=https://www.sciencedirect.com/science/article/abs/pii/S1439179117304498|journal=Basic and Applied Ecology|language=en|volume=31|pages=1–9|doi=10.1016/j.baae.2018.05.009|issn=1439-1791|last1=Walas|first1=Łukasz|last2=Mandryk|first2=Wojciech|last3=Thomas|first3=Peter A.|last4=Tyrała-Wierucka|first4=Żanna|last5=Iszkuło|first5=Grzegorz|s2cid=90740232}}</ref> but almost all conifers are monoecious.<ref name="Walas">{{cite journal| vauthors = Walas Ł, Mandryk W, Thomas PA, Tyrała-Wierucka Ż, Iszkuło G |date=2018|title=Sexual systems in gymnosperms: A review|journal=Basic and Applied Ecology|volume=31|pages=1–9|doi=10.1016/j.baae.2018.05.009|s2cid=90740232|url=http://eprints.keele.ac.uk/4961/1/29052018_1-s2.0-S1439179117304498-main.pdf}}</ref>
In gymnosperms, the sexual systems dioecy and monoecy are strongly correlated with the mode of pollen dispersal, monoecious species are predominantly wind dispersed ([[Biology:Anemophily|anemophily]]) and dioecious species animal-dispersed ([[Biology:Zoophily|zoophily]]).<ref name=Givnish>{{cite journal |author=Givnish, TJ |date=1980 |title=Ecological constraints on the evolution of breeding systems in seed plants: dioecy and dispersal in gymnosperms |journal=Evolution |volume=34 |issue=5 |pages=959–972 |doi=10.1111/j.1558-5646.1980.tb04034.x |pmid=28581147 |doi-access=free }}</ref>

About 6 percent of [[Biology:Angiosperm|flowering plant]] species are entirely dioecious and about 7% of angiosperm [[Biology:Genus|genera]] contain some dioecious species.<ref name="Renner & Ricklefs">{{cite journal|author1=Renner, S. S. |author2=R. E. Ricklefs|year=1995|title=Dioecy and its correlates in the flowering plants|journal=American Journal of Botany|volume=82|issue=5|pages=596–606|doi=10.2307/2445418|jstor=2445418|url=https://epub.ub.uni-muenchen.de/14619/}}</ref> Dioecy is more common in [[Biology:Woody plant|woody plant]]s,<ref>{{citation |author1=Matallana, G. |author2=Wendt, T. |author3=Araujo, D.S.D. |author4=Scarano, F.R. |year=2005 |title=High abundance of dioecious plants in a tropical coastal vegetation |journal=American Journal of Botany |volume=92 |issue=9 |pages=1513–1519 |doi=10.3732/ajb.92.9.1513 |pmid=21646169 }}</ref> and heterotrophic species.<ref>{{cite journal | author = Nickrent D.L., Musselman L.J. | year = 2004 | title = Introduction to Parasitic Flowering Plants | url = http://www.apsnet.org/edcenter/intropp/pathogengroups/pages/parasiticplants.aspx | journal = The Plant Health Instructor | doi = 10.1094/PHI-I-2004-0330-01 | access-date = 2017-01-10 | archive-url = https://web.archive.org/web/20161005194620/http://www.apsnet.org/edcenter/intropp/pathogengroups/pages/parasiticplants.aspx | archive-date = 2016-10-05 | url-status = dead }}</ref> 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 ''[[Biology:Arisaema|Arisaema]]'' species.<ref>{{Cite book|last1=Fusco|first1=Giuseppe|url=https://books.google.com/books?id=AKGsDwAAQBAJ&q=gonochorism+sex+determination|title=The Biology of Reproduction|last2=Minelli|first2=Alessandro|date=2019-10-10|publisher=Cambridge University Press|isbn=978-1-108-49985-9|pages=329|language=en}}</ref>

Certain [[Biology:Algae|algae]] are dioecious.{{clarify|reason=specify which groups and provide some examples|date=August 2021}}<ref>Maggs, C.A. and Hommersand, M.H. 1993. ''Seaweeds of the British Isles Volume 1 Rhodophyta Part 3A Ceramiales''. The Natural History Museum, London. {{ISBN|0-11-310045-0}}</ref> Dioecy is prevalent in the brown algae ([[Biology:Phaeophyceae|Phaeophyceae]]) and may have been the ancestral state in that group.<ref name=Luthringer>{{cite journal |vauthors=LuthringerR, Cormier A, Ahmed S, Peters AF, Cock JM, Coelho, SM |title=Sexual dimorphism in the brown algae |journal=Perspectives in Phycology |volume=1 |date=2014 |issue=1 |pages=11–25 |doi=10.1127/2198-011X/2014/0002 }}</ref>

=== Evolution of dioecy ===
In plants, dioecy has [[Biology:Convergent evolution|evolved independently]] multiple times<ref>{{Cite journal |last1=Bachtrog |first1=Doris |last2=Mank |first2=Judith E. |last3=Peichel |first3=Catherine L. |last4=Kirkpatrick |first4=Mark |last5=Otto |first5=Sarah P. |last6=Ashman |first6=Tia-Lynn |last7=Hahn |first7=Matthew W. |last8=Kitano |first8=Jun |last9=Mayrose |first9=Itay |last10=Ming |first10=Ray |last11=Perrin |first11=Nicolas |date=2014-07-01 |title=Sex Determination: Why So Many Ways of Doing It? |journal=PLOS Biology |volume=12 |issue=7 |pages=e1001899 |doi=10.1371/journal.pbio.1001899 |issn=1544-9173 |pmc=4077654 |pmid=24983465 |doi-access=free }}</ref> generally either from hermaphroditic species or from monoecious species. A previously untested hypothesis is that this reduces inbreeding;<ref>{{Cite journal |last1=Sarkar |first1=Sutanu |last2=Banerjee |first2=Joydeep |last3=Gantait |first3=Saikat|date=2017-05-29 |title=Sex-oriented research on dioecious crops of Indian subcontinent: an updated review|url=https://doi.org/10.1007/s13205-017-0723-8|journal=3 Biotech|language=en|volume=7|issue=2|pages=93|doi=10.1007/s13205-017-0723-8|issn=2190-5738|pmc=5447520|pmid=28555429}}</ref> dioecy has been shown to be associated with increased [[Biology:Genetic diversity|genetic diversity]] and greater protection against deleterious mutations.<ref name="Muyle">{{Cite journal |last1=Muyle|first1=Aline|last2=Martin |first2=Hélène|last3=Zemp|first3=Niklaus|last4=Mollion|first4=Maéva |last5=Gallina|first5=Sophie|last6=Tavares|first6=Raquel |last7=Silva|first7=Alexandre|last8=Bataillon|first8=Thomas|last9=Widmer |first9=Alex|last10=Glémin|first10=Sylvain |last11=Touzet|first11=Pascal |date=2021-03-01 |title=Dioecy Is Associated with High Genetic Diversity and Adaptation Rates in the Plant Genus Silene|url=https://doi.org/10.1093/molbev/msaa229 |journal=Molecular Biology and Evolution |volume=38 |issue=3 |pages=805–818|doi=10.1093/molbev/msaa229|issn=0737-4038|pmc=7947750|pmid=32926156}}</ref>  Regardless of the evolutionary pathway the intermediate states need to have fitness advantages compared to  cosexual flowers in order to survive.<ref>{{Cite book|last=Cruzan|first=Mitchell B.|url=https://books.google.com/books?id=nzRtDwAAQBAJ&q=Dioicy+sexual+system&pg=PA377|title=Evolutionary Biology: A Plant Perspective|date=2018-09-11|publisher=Oxford University Press|isbn=978-0-19-088268-6|pages=377|language=en}}</ref>

Dioecy evolves due to male or female sterility,<ref>{{Cite book|last1=Atwell|first1=Brian James|url=https://books.google.com/books?id=chWs4ewSzpEC&q=all+botanists+agree+dioecy&pg=PT49|title=Plants in Action: Adaptation in Nature, Performance in Cultivation|last2=Kriedemann|first2=Paul E.|last3=Turnbull|first3=Colin G. N.|date=1999|publisher=Macmillan Education AU|isbn=978-0-7329-4439-1|pages=249|language=en}}</ref> although it is unlikely that mutations for male and female sterility occurred at the same time.<ref>{{Cite book|last=Karasawa|first=Marines Marli Gniech|url=https://books.google.com/books?id=prsDCwAAQBAJ&q=evolution+Monoecy|title=Reproductive Diversity of Plants: An Evolutionary Perspective and Genetic Basis|date=2015-11-23|publisher=Springer|isbn=978-3-319-21254-8|pages=31|language=en}}</ref> In angiosperms unisexual flowers evolve from bisexual ones.<ref>{{Cite book|last1=Núñez-Farfán|first1=Juan|url=https://books.google.com/books?id=iF70DwAAQBAJ&q=dioecy+consensus&pg=PA177|title=Evolutionary Ecology of Plant-Herbivore Interaction|last2=Valverde|first2=Pedro Luis|date=2020-07-30|publisher=Springer Nature|isbn=978-3-030-46012-9|pages=177|language=en}}</ref> Dioecy occurs in almost half of plant families, but only in a minority of genera, suggesting recent evolution.<ref>{{Cite book|last=Reeve|first=Eric C. R.|url=https://books.google.com/books?id=PuCYAgAAQBAJ&q=evolution+of+dioecy&pg=PA616|title=Encyclopedia of Genetics|date=2014-01-14|publisher=Routledge|isbn=978-1-134-26350-9|pages=616|language=en}}</ref> For 160 families that have dioecious species, dioecy is thought to have evolved more than 100 times.<ref>{{Cite journal|last=Ainsworth|first=Charles|date=2000-08-01|title=Boys and Girls Come Out to Play: The Molecular Biology of Dioecious Plants|journal=Annals of Botany|volume=86|issue=2|pages=211–221|doi=10.1006/anbo.2000.1201|s2cid=85039623 |issn=0305-7364|doi-access=free}}</ref>

In the family [[Biology:Caricaceae|Caricaceae]], dioecy is likely the ancestral sexual system.<ref>{{Cite book|last=Mitra|first=Sisir|url=https://books.google.com/books?id=7zX8DwAAQBAJ&dq=trioecy&pg=PA161|title=The Papaya: Botany, Production and Uses|date=2020-09-01|publisher=CABI|isbn=978-1-78924-190-7|pages=161|language=en}}</ref>

==== From monoecy ====
Dioecious flowering plants can evolve from monoecious ancestors that have flowers containing both functional stamens and functional carpels.<ref name="Bawa80" />  Some authors argue monoecy and dioecy are related.<ref name="Batygina">{{Cite book|last=Batygina|first=T. B.|url=https://books.google.com/books?id=4VOWDwAAQBAJ&q=monoecy&pg=PA43|title=Embryology of Flowering Plants: Terminology and Concepts, Vol. 3: Reproductive Systems|date=2019-04-23|publisher=CRC Press|isbn=978-1-4398-4436-6|pages=43|language=en}}</ref>

In the genus ''[[Biology:Sagittaria|Sagittaria]]'', since there is a distribution of sexual systems, it has been postulated that dioecy evolved from monoecy<ref>{{Cite book|last1=Wilson|first1=Karen L.|url=https://books.google.com/books?id=YzQBUQqLS0YC&q=Monoecy&pg=PA264|title=Monocots: Systematics and Evolution: Systematics and Evolution|last2=Morrison|first2=David A.|date=2000-05-19|publisher=Csiro Publishing|isbn=978-0-643-09929-6|pages=264|language=en}}</ref> through [[Biology:Gynodioecy|gynodioecy]] mainly from mutations that resulted in male sterility.<ref name="encyclobiol">{{Cite book|url=https://books.google.com/books?id=_r4OCAAAQBAJ&q=evolution+of+Monoecy&pg=RA1-PA478|title=Encyclopedia of Evolutionary Biology|date=2016-04-14|publisher=Academic Press|isbn=978-0-12-800426-5|volume=2|language=en}}</ref>{{rp|478}} However, since the ancestral state is unclear, more work is needed to clarify the evolution of dioecy via monoecy.<ref name="encyclobiol" />{{rp|478}}

==== From hermaphroditism ====
Dioecy usually evolves from [[Biology:Hermaphrodite#Plants|hermaphroditism]] through [[Biology:Gynodioecy|gynodioecy]] but may also evolve through [[Biology:Androdioecy|androdioecy]],<ref>{{Cite journal|last1=Perry|first1=Laura E.|last2=Pannell|first2=John R.|last3=Dorken|first3=Marcel E.|date=2012-04-19|title=Two's Company, Three's a Crowd: Experimental Evaluation of the Evolutionary Maintenance of Trioecy in Mercurialis annua (Euphorbiaceae)|journal=PLOS ONE|language=en|volume=7|issue=4|pages=e35597|doi=10.1371/journal.pone.0035597|issn=1932-6203|pmc=3330815|pmid=22532862|bibcode=2012PLoSO...735597P |doi-access=free}}</ref> through [[Biology:Distyly|distyly]]<ref>{{Cite book|last=Leonard|first=Janet L.|url=https://books.google.com/books?id=0rWZDwAAQBAJ&q=evolution+of+Monoecy&pg=PA91|title=Transitions Between Sexual Systems: Understanding the Mechanisms of, and Pathways Between, Dioecy, Hermaphroditism and Other Sexual Systems|date=2019-05-21|publisher=Springer|isbn=978-3-319-94139-4|pages=91|language=en}}</ref> or through [[Biology:Heterostyly|heterostyly]].<ref name="Muyle"/> In the [[Biology:Asteraceae|Asteraceae]], dioecy may have [[Biology:Convergent evolution|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.<ref name="Landry">{{Cite book|last1=Landry|first1=Christian R.|url=https://books.google.com/books?id=WATFBAAAQBAJ&q=Asteraceae+dioecy&pg=PA9|title=Ecological Genomics: Ecology and the Evolution of Genes and Genomes|last2=Aubin-Horth|first2=Nadia|date=2013-11-25|publisher=Springer Science & Business Media|isbn=978-94-007-7347-9|pages=9|language=en}}</ref>

In ''[[Biology:Silene|Silene]]'', since there is no monoecy, it is suggested that dioecy evolved through gynodioecy.<ref>{{Cite journal|last1=Casimiro-Soriguer|first1=Inés|last2=Buide|first2=Maria L.|last3=Narbona|first3=Eduardo|date=2015-01-01|title=Diversity of sexual systems within different lineages of the genus Silene|url=https://doi.org/10.1093/aobpla/plv037|journal=AoB Plants|volume=7|issue=plv037|pages=plv037|doi=10.1093/aobpla/plv037|pmid=25862920|issn=2041-2851|pmc=4433491}}</ref>

==In mycology==
Very few dioecious fungi have been discovered.<ref>{{Cite book|last=Gupta|first=Rajni|url=https://books.google.com/books?id=uPCATfIDZBoC&q=dioecy+in+fungi&pg=PA77|title=A Textbook of Fungi|publisher=APH Publishing|isbn=978-81-7648-737-5|pages=77|language=en}}</ref>

Monoecy and dioecy in [[Biology:Fungus|fungi]] 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 [[Biology:Karyogamy|karyogamy]] to form a [[Biology:Zygote|zygote]].<ref name="Esser">{{cite journal |author=Esser, K. |year=1971 |title=Breeding systems in fungi and their significance for genetic recombination |journal=Molecular and General Genetics |volume=110 |issue=1 |pages=86–100 |doi=10.1007/bf00276051|pmid=5102399 |s2cid=11353336 }}</ref> The definition avoids reference to male and female reproductive structures, which are rare in fungi.<ref name="Esser" /> 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.<ref name="Esser" />

==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 [[Biology:Allogamy|allogamy]] (outcrossing), and thus tends to reduce the expression of recessive deleterious mutations present in a population.<ref name=Charlesworth>{{cite journal |vauthors=Charlesworth D, Willis JH |title=The genetics of inbreeding depression |journal=Nat. Rev. Genet. |volume=10 |issue=11 |pages=783–96 |year=2009 |pmid=19834483 |doi=10.1038/nrg2664 |s2cid=771357 }}</ref> 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.<ref name="BruijningVisser2017">{{cite journal |last1=Bruijning |first1=Marjolein |last2=Visser |first2=Marco D. |last3=Muller-Landau |first3=Helene C. |last4=Wright |first4=S. Joseph |last5=Comita |first5=Liza S. |last6=Hubbell |first6=Stephen P. |last7=de Kroon |first7=Hans |last8=Jongejans |first8=Eelke |title=Surviving in a Cosexual World: A Cost-Benefit Analysis of Dioecy in Tropical Trees |journal=The American Naturalist |volume=189 |issue=3 |year=2017 |pages=297–314 |issn=0003-0147 |doi=10.1086/690137|pmid=28221824 |hdl=2066/168955 |s2cid=6839285 |hdl-access=free }}</ref> Dioecy may also accelerate or retard lineage diversification in [[Biology:Flowering plant|angiosperms]]. 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.<ref name="SabathGoldberg2016">{{cite journal |last1=Sabath |first1=Niv |last2=Goldberg |first2=Emma E. |last3=Glick |first3=Lior |last4=Einhorn |first4=Moshe |last5=Ashman |first5=Tia-Lynn |last6=Ming |first6=Ray |last7=Otto |first7=Sarah P. |last8=Vamosi |first8=Jana C. |last9=Mayrose |first9=Itay |title=Dioecy does not consistently accelerate or slow lineage diversification across multiple genera of angiosperms |journal=New Phytologist |volume=209 |issue=3 |year=2016 |pages=1290–1300 |doi=10.1111/nph.13696|pmid=26467174 |doi-access=free }}</ref>

== See also ==
*[[Biology:Gonochorism|Gonochorism]]
*[[Biology:Hermaphrodite|Hermaphrodite]]
*[[Biology:Plant reproductive morphology|Plant reproductive morphology]]
*Self-incompatibility in plants
*[[Biology:Sexual dimorphism|Sexual dimorphism]]
*[[Biology:Trioecy|Trioecy]]

== References ==
{{reflist|refs=
<ref name="Kliman-2016">{{cite book |last=Kliman |first=Richard |date=2016 |title=Encyclopedia of Evolutionary Biology |publisher=Academic Press |isbn=978-0-12-800426-5 |url=https://books.google.com/books?id=_r4OCAAAQBAJ |archive-url=https://web.archive.org/web/20210506205920/https://www.google.com/books/edition/Encyclopedia_of_Evolutionary_Biology/_r4OCAAAQBAJ?hl=en&gbpv=0&kptab=overview |archive-date=May 6, 2021}} [https://archive.org/details/encyclopediaofevolutionarybiology Alternative archive URL]</ref>

<ref name=Bawa80>{{cite journal|title=Evolution of Dioecy in Flowering Plants|author=K. S. Bawa |journal=Annual Review of Ecology and Systematics|volume=11|year=1980 |pages=15–39 |doi=10.1146/annurev.es.11.110180.000311|jstor = 2096901}}</ref> 
}}

==Bibliography==
*{{Cite book |last=Beentje |first=Henk |year=2010 |title=The Kew Plant Glossary |publication-place=Richmond, Surrey |publisher=[[Organization:Royal Botanic Gardens, Kew|Royal Botanic Gardens, Kew]] |isbn=978-1-84246-422-9 }}
*{{Cite book |last=Mauseth |first=James D. |title=Botany : An Introduction to Plant Biology |edition=5th |year=2014 |isbn=978-1-4496-6580-7 |publisher=Jones and Bartlett Learning |location=Sudbury, MA}}

[[Category:Plant sexuality]]
[[Category:Dioecious plants|*]]
[[Category:Sexual reproduction]]
[[Category:Reproductive system]]

{{Sourceattribution|Dioecy}}