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{{short description|System of plant reproduction}}
'''Distyly''' is a breeding system in plants that is characterized by two separate flower morphs, where individual plants produce flowers that have either long styles and short stamens (L-morph flowers) or short styles and long stamens (S-morph flowers).<ref>{{cite journal| vauthors = Lewis D |date=1942|title=The Physiology of Incompatibility in Plants. I. The Effect of Temperature|journal=Proceedings of the Royal Society of London. Series B, Biological Sciences|volume=131|issue=862|pages=13–26|doi=10.1098/rspb.1942.0015|jstor=82364|bibcode=1942RSPSB.131...13L|s2cid=84753102|issn=0080-4649|doi-access=}}</ref> However, distyly can refer to any plant that shows some degree of [[Self-incompatibility#/Sporophytic self-incompatibility SSI|self-incompatibility]] and has two morphs if at least one of the following characteristics is true; there is a difference in [[Stigma (botany)#/Style|style]] length, [[Stamen#/Morphology and terminology|filament]] length, [[pollen]] size or shape, or the surface of the [[Stigma (botany)|stigma]].<ref>{{cite journal| vauthors = Muenchow G |date=August 1982|title=A loss-of-alleles model for the evolution of distyly |journal=Heredity|language=en|volume=49|issue=1|pages=81–93|doi=10.1038/hdy.1982.67|issn=0018-067X|doi-access=free|bibcode=1982Hered..49...81M }}</ref> Specifically these plants exhibit intra-morph self-incompatibility, flowers of the same style morph are incompatible.<ref>{{cite book |vauthors = Barrett SC, Cruzan MB | series = Advances in Cellular and Molecular Biology of Plants| chapter = Incompatibility in heterostylous plants|date=1994| title = Genetic control of self-incompatibility and reproductive development in flowering plants| volume = 2|pages=189–219|publisher=Springer Netherlands|isbn=978-90-481-4340-5|doi=10.1007/978-94-017-1669-7_10}}</ref> Distylous species that do not exhibit true self-incompatibility generally show a bias towards inter-morph crosses - meaning they exhibit higher success rates when reproducing with an individual of the opposite morph.<ref>{{cite journal | vauthors = Shao JW, Wang HF, Fang SP, Conti E, Chen YJ, Zhu HM | title = Intraspecific variation of self-incompatibility in the distylous plant ''Primula merrilliana'' | journal = AoB Plants | volume = 11 | issue = 3 | article-number = plz030 | date = June 2019 | pmid = 32489575 | pmc = 6557196 | doi = 10.1093/aobpla/plz030 }}</ref>

Distyly is a type of [[heterostyly]] in which a plant demonstrates [[Herkogamy|reciprocal herkogamy]].

[[File:Distyly.png|thumb|279x279px|Diagram of both distylous morphs]]
[[File:Distyly primula.jpg|thumb|312x312px|Example of distyly in Primula. A. L-morph (pin), B. S-morph (thrum)

1. petal. 2 sepal. 3 anther. 4 pistil. ]]

== Background ==
The first scientific account of distyly can be found in Stephan Bejthe's Caroli book ''Clusii Atrebatis Rariorum aliquot stirpium'' <ref>{{Cite book |last=Bejthe |first=Stephan |url=https://www.biodiversitylibrary.org/bibliography/845 |title=Caroli Clusii Atrebatis Rariorum aliquot stirpium :per Pannoniam, Austriam, & vicinas quasdam provincias observatarum historia, quatuor libris expressa |date=1583 |publisher=Ex officina Christophori Plantini |location=Antverpiae |doi=10.5962/bhl.title.845}}</ref>''.'' Bejthe describes the two floral morphs of ''Primula veris.'' [[Charles Darwin]] popularized distyly with his account of it in his book ''[[The Different Forms of Flowers on Plants of the Same Species]]''.<ref name=":0">{{cite book |title=The different forms of flowers on plants of the same species by Charles Darwin ... |vauthors=Darwin C |publisher=D. Appleton and Co |year=1877 |oclc=894148387}}</ref> Darwin's book represents the first account of intramorphic self-incompatibility in distylous plants and focuses on garden experiments in which he looks at seed set of different distylous ''Primula''. Darwin names the two floral morphs S- and L-morph, moving away from the [[vernacular]] names, Pin (for L-morph) and Thrum (for S-morph), which he states were initially assigned by florist.

Distylous species have been identified in 28 families of Angiosperm, likely evolving independently in each family.<ref name=":2" /> This means, the system has evolved at least 28 times, though it has been suggested the system has evolved multiple times within some families.<ref name=":2" /> Since distyly has evolved more than once, it is considered a case of [[convergent evolution]].<ref name=":2" />

== Reciprocal herkogamy ==
Reciprocal herkogamy likely evolved to prevent the pollen of the same flower from landing on its own stigma. This in turn promotes outcrossing.

In a study of ''Primula veris'' it was found that pin flowers exhibit higher rates of self-pollination and capture more pollen than the thrum morph.<ref name=":3">{{cite journal| vauthors = Naiki A |date=2012|title=Heterostyly and the possibility of its breakdown by polyploidization|journal=Plant Species Biology|language=en|volume=27|issue=1 |pages=3–29|doi=10.1111/j.1442-1984.2011.00363.x|bibcode=2012PSBio..27....3N }}</ref> Different [[pollinator]]s show varying levels of success while pollinating the different ''Primula'' morphs, the head or proboscis length of a pollinator is positively correlated to the uptake of pollen from long styled flowers and negatively correlated for pollen uptake on short styled flowers.<ref name=":1" /> The opposite is true for pollinators with smaller heads, such as bees, they uptake more pollen from short styled morphs than long styled ones.<ref name=":1">{{cite journal| vauthors = Deschepper P, Brys R, Jacquemyn H |date=2018-03-01|title=The impact of flower morphology and pollinator community composition on pollen transfer in the distylous Primula veris|journal=Botanical Journal of the Linnean Society|volume=186|issue=3|pages=414–424|doi=10.1093/botlinnean/box097|issn=0024-4074}}</ref> The differentiation in pollinators allows the plants to reduce levels of intra-morph pollination.

== Models of evolution ==
There are two main hypothetical models for the order in which the traits of distyly evolved, the 'selfing avoidance model' <ref name=":6">{{cite journal| vauthors = Charlesworth D, Charlesworth B |date=October 1979|title=A Model for the Evolution of Distyly |journal=The American Naturalist|language=en|volume=114|issue=4|pages=467–498|doi=10.1086/283496|bibcode=1979ANat..114..467C |s2cid=85285185|issn=0003-0147}}</ref> and the 'pollen transfer model'.<ref name=":7">{{cite book | vauthors = Lloyd DG, Webb CJ | chapter = The Selection of Heterostyly|date=1992 |title = Evolution and Function of Heterostyly| series = Monographs on Theoretical and Applied Genetics|volume=15|pages=179–207| veditors = Barrett SC |place=Berlin, Heidelberg|publisher=Springer Berlin Heidelberg|doi=10.1007/978-3-642-86656-2_7|isbn=978-3-642-86658-6 }}</ref>

# The selfing avoidance model suggests self-incompatibility (SI) evolved first, followed by the morphological difference. It was suggested that the male component of SI would evolve first via a recessive mutation, followed by female characteristics via a dominant mutation, and finally male morphological differences would evolve via a third mutation.<ref name=":6" />
# The pollen transfer model argues that morphological differences evolved first, and if a species is facing [[inbreeding depression]], it may evolve SI.<ref name=":7" /> This model can be used to explain the presence of reciprocal [[herkogamy]] in self-compatible species.<ref name=":2" />

== Genetic control of distyly ==

A [[supergene]], called the self-incompatibility (or S''-'') locus, is responsible for the occurrence of distyly.<ref name=":2">{{cite journal | vauthors = Barrett SC | title = 'A most complex marriage arrangement': recent advances on heterostyly and unresolved questions | journal = The New Phytologist | volume = 224 | issue = 3 | pages = 1051–1067 | date = November 2019 | pmid = 31631362 | doi = 10.1111/nph.16026 | doi-access = free | bibcode = 2019NewPh.224.1051B }}</ref> The ''S''-locus is composed of three tightly linked genes (''S''-genes) which [[Mendelian inheritance|segregate]] as a single unit.<ref name=":2" />

Traditionally it was hypothesized that one ''S-''gene controls all female aspects of distyly, one gene that controls the male morphological aspects, and one gene that determines the male [[mating type]].<ref>{{cite journal | vauthors = Kappel C, Huu CN, Lenhard M | title = A short story gets longer: recent insights into the molecular basis of heterostyly | journal = Journal of Experimental Botany | volume = 68 | issue = 21–22 | pages = 5719–5730 | date = December 2017 | pmid = 29099983 | doi = 10.1093/jxb/erx387 | doi-access = free }}</ref> While this hypothesis appears to be true in ''[[Turnera]]'',<ref name=":4" /> it is not true in ''[[Primula]]'' <ref name="hull-repository.worktribe.com">{{cite journal | vauthors = Li J, Cocker JM, Wright J, Webster MA, McMullan M, Dyer S, Swarbreck D, Caccamo M, Oosterhout CV, Gilmartin PM | display-authors = 6 | title = Genetic architecture and evolution of the S locus supergene in Primula vulgaris | journal = Nature Plants | volume = 2 | issue = 12 | page = 16188 | date = December 2016 | pmid = 27909301 | doi = 10.1038/nplants.2016.188 | bibcode = 2016NatPl...216188L | s2cid = 205458474 | url = https://hull-repository.worktribe.com/output/3712561 }}</ref> nor ''[[Linum]]''.<ref name=":12">{{Cite journal |last1=Gutiérrez-Valencia |first1=Juanita |last2=Fracassetti |first2=Marco |last3=Berdan |first3=Emma L. |last4=Bunikis |first4=Ignas |last5=Soler |first5=Lucile |last6=Dainat |first6=Jacques |last7=Kutschera |first7=Verena E. |last8=Losvik |first8=Aleksandra |last9=Désamoré |first9=Aurélie |last10=Hughes |first10=P. William |last11=Foroozani |first11=Alireza |last12=Laenen |first12=Benjamin |last13=Pesquet |first13=Edouard |last14=Abdelaziz |first14=Mohamed |last15=Pettersson |first15=Olga Vinnere |date=2022 |title=Genomic analyses of the Linum distyly supergene reveal convergent evolution at the molecular level |journal=Current Biology |language=en |volume=32 |issue=20 |pages=4360–4371.e6 |doi=10.1016/j.cub.2022.08.042|pmid=36087578 |s2cid=249242025 |doi-access=free |bibcode=2022CBio...32E4360G |hdl=10481/78952 |hdl-access=free }}</ref> The S-morph is [[hemizygous]] for the ''S-''locus and the L-morph does not have an [[Allele|allelic]] counterpart <ref name=":2" />''.'' The hemizygotic nature of the ''S-''locus has been shown in ''Primula'' <ref name="hull-repository.worktribe.com" /> '','' ''[[Gelsemium]]'',<ref name=":16">{{Cite journal |last1=Zhao |first1=Zhongtao |last2=Zhang |first2=Yu |last3=Shi |first3=Miaomiao |last4=Liu |first4=Zhaoying |last5=Xu |first5=Yuanqing |last6=Luo |first6=Zhonglai |last7=Yuan |first7=Shuai |last8=Tu |first8=Tieyao |last9=Sun |first9=Zhiliang |last10=Zhang |first10=Dianxiang |last11=Barrett |first11=Spencer C. H. |date=2022-11-22 |title=Genomic evidence supports the genetic convergence of a supergene controlling the distylous floral syndrome |url=https://onlinelibrary.wiley.com/doi/10.1111/nph.18540 |journal=New Phytologist |volume=237 |issue=2 |language=en |pages=601–614 |doi=10.1111/nph.18540 |pmid=36239093 |s2cid=252897518 |issn=0028-646X|url-access=subscription }}</ref> ''Linum'' <ref name=":13">{{cite journal | vauthors = Ushijima K, Nakano R, Bando M, Shigezane Y, Ikeda K, Namba Y, Kume S, Kitabata T, Mori H, Kubo Y | display-authors = 6 | title = Isolation of the floral morph-related genes in heterostylous flax (Linum grandiflorum): the genetic polymorphism and the transcriptional and post-transcriptional regulations of the S locus | journal = The Plant Journal | volume = 69 | issue = 2 | pages = 317–31 | date = January 2012 | pmid = 21923744 | doi = 10.1111/j.1365-313X.2011.04792.x | doi-access = }}</ref><ref name=":12" />'', [[Fagopyrum]]'' <ref name=":14">{{cite journal | vauthors = Yasui Y, Mori M, Aii J, Abe T, Matsumoto D, Sato S, Hayashi Y, Ohnishi O, Ota T | display-authors = 6 | title = S-LOCUS EARLY FLOWERING 3 is exclusively present in the genomes of short-styled buckwheat plants that exhibit heteromorphic self-incompatibility | journal = PLOS ONE | volume = 7 | issue = 2 | article-number = e31264 | date = 2012-02-01 | pmid = 22312442 | pmc = 3270035 | doi = 10.1371/journal.pone.0031264 | bibcode = 2012PLoSO...731264Y | doi-access = free }}</ref><ref name=":17">{{Cite journal |last1=Fawcett |first1=Jeffrey A. |last2=Takeshima |first2=Ryoma |last3=Kikuchi |first3=Shinji |last4=Yazaki |first4=Euki |last5=Katsube-Tanaka |first5=Tomoyuki |last6=Dong |first6=Yumei |last7=Li |first7=Meifang |last8=Hunt |first8=Harriet V. |last9=Jones |first9=Martin K. |last10=Lister |first10=Diane L. |last11=Ohsako |first11=Takanori |last12=Ogiso-Tanaka |first12=Eri |last13=Fujii |first13=Kenichiro |last14=Hara |first14=Takashi |last15=Matsui |first15=Katsuhiro |year=2023 |title=Genome sequencing reveals the genetic architecture of heterostyly and domestication history of common buckwheat |url=https://www.nature.com/articles/s41477-023-01474-1 |journal=Nature Plants |language=en |volume=9 |issue=8 |pages=1236–1251 |doi=10.1038/s41477-023-01474-1 |pmid=37563460 |bibcode=2023NatPl...9.1236F |issn=2055-0278|url-access=subscription }}</ref>'','' ''Turnera,''<ref name=":4">{{cite journal | vauthors = Shore JS, Hamam HJ, Chafe PD, Labonne JD, Henning PM, McCubbin AG | title = The long and short of the S-locus in Turnera (Passifloraceae) | journal = The New Phytologist | volume = 224 | issue = 3 | pages = 1316–1329 | date = November 2019 | pmid = 31144315 | doi = 10.1111/nph.15970 | doi-access = free | bibcode = 2019NewPh.224.1316S }}</ref> ''[[Nymphoides]]''<ref name=":18">{{Cite journal |last1=Yang |first1=Jingshan |last2=Xue |first2=Haoran |last3=Li |first3=Zhizhong |last4=Zhang |first4=Yue |last5=Shi |first5=Tao |last6=He |first6=Xiangyan |last7=Barrett |first7=Spencer C. H. |last8=Wang |first8=Qingfeng |last9=Chen |first9=Jinming |date=2023-09-17 |title=Haplotype-resolved genome assembly provides insights into the evolution of S -locus supergene in distylous Nymphoides indica |journal=New Phytologist |volume=240 |issue=5 |pages=2058–2071 |language=en |doi=10.1111/nph.19264 |issn=0028-646X|doi-access=free |pmid=37717220 |bibcode=2023NewPh.240.2058Y }}</ref> and ''[[Chrysojasminum]]''.<ref name=":21">{{Cite journal |last1=Raimondeau |first1=Pauline |last2=Ksouda |first2=Sayam |last3=Marande |first3=William |last4=Fuchs |first4=Anne-Laure |last5=Gryta |first5=Hervé |last6=Theron |first6=Anthony |last7=Puyoou |first7=Aurore |last8=Dupin |first8=Julia |last9=Cheptou |first9=Pierre-Olivier |last10=Vautrin |first10=Sonia |last11=Valière |first11=Sophie |last12=Manzi |first12=Sophie |last13=Baali-Cherif |first13=Djamel |last14=Chave |first14=Jérôme |last15=Christin |first15=Pascal-Antoine |date=May 2024 |title=A hemizygous supergene controls homomorphic and heteromorphic self-incompatibility systems in Oleaceae |journal=Current Biology |volume=34 |issue=9 |pages=1977–1986.e8 |doi=10.1016/j.cub.2024.03.029 |pmid=38626764 |bibcode=2024CBio...34.1977R |issn=0960-9822}}</ref>

The presence of the ''S-''locus results in changes to [[gene expression]] between the two floral morphs, as has been demonstrated using [[Transcriptomics technologies|transcriptomic analyses]] of ''[[Lithospermum|Lithospermum multiflorum]]'' <ref>{{cite journal | vauthors = Cohen JI | title = ''De novo'' Sequencing and Comparative Transcriptomics of Floral Development of the Distylous Species ''Lithospermum multiflorum'' | journal = Frontiers in Plant Science | volume = 7 | page = 1934 | date = 2016-12-23 | pmid = 28066486 | pmc = 5179544 | doi = 10.3389/fpls.2016.01934 | doi-access = free }}</ref> '', [[Primula veris]]'',<ref name=":5">{{cite journal |vauthors=Nowak MD, Russo G, Schlapbach R, Huu CN, Lenhard M, Conti E |date=January 2015 |title=The draft genome of Primula veris yields insights into the molecular basis of heterostyly |journal=Genome Biology |volume=16 |issue=1 |page=12 |doi=10.1186/s13059-014-0567-z |pmc=4305239 |pmid=25651398 |doi-access=free }}</ref> ''[[Primula|Primula oreodoxa]]'' <ref>{{cite journal | vauthors = Zhao Z, Luo Z, Yuan S, Mei L, Zhang D | title = Global transcriptome and gene co-expression network analyses on the development of distyly in Primula oreodoxa | journal = Heredity | volume = 123 | issue = 6 | pages = 784–794 | date = December 2019 | pmid = 31308492 | pmc = 6834660 | doi = 10.1038/s41437-019-0250-y | bibcode = 2019Hered.123..784Z }}</ref>'', [[Primula vulgaris]]'' <ref>{{cite journal | vauthors = Burrows B, McCubbin A | title = Examination of ''S''-Locus Regulated Differential Expression in ''Primula vulgaris'' Floral Development | journal = Plants | volume = 7 | issue = 2 | page = 38 | date = May 2018 | pmid = 29724049 | pmc = 6027539 | doi = 10.3390/plants7020038 | doi-access = free | bibcode = 2018Plnts...7...38B }}</ref> and ''[[Turnera subulata]]<ref>{{cite journal |vauthors=Henning PM, Shore JS, McCubbin AG |date=June 2020 |title=Transcriptome and Network Analyses of Heterostyly in ''Turnera subulata'' Provide Mechanistic Insights: Are ''S''-Loci a Red-Light for Pistil Elongation? |journal=Plants |volume=9 |issue=6 |page=713 |doi=10.3390/plants9060713 |pmc=7356734 |pmid=32503265 |doi-access=free|bibcode=2020Plnts...9..713H }}</ref>'', and ''[[Forsythia suspensa]]''.<ref name=":20">{{Cite journal |last1=Song |first1=Yun |last2=Li |first2=Zheng |last3=Du |first3=Xiaorong |last4=Li |first4=Aoxuan |last5=Cao |first5=Yaping |last6=Jia |first6=Mengjun |last7=Niu |first7=Yanbing |last8=Qiao |first8=Yonggang |date=2024-05-06 |title=Study on the brassinosteroids modulated regulation of the style growth in Forsythia suspensa (Thunb.) Vahl |url=https://link.springer.com/10.1007/s10725-024-01149-7 |journal=Plant Growth Regulation |volume=103 |issue=3 |pages=763–774 |language=en |doi=10.1007/s10725-024-01149-7 |bibcode=2024PGroR.103..763S |issn=0167-6903|url-access=subscription }}</ref>

=== The ''S''-locus of ''Chrysojasminum'' ===
In ''Chrysojasminum'', the ''S''-locus is composed of two ''S''-genes, ''BZR1'' and ''GA2ox''.<ref name=":21" /> GA2ox is hypothetically involved in establishing self-incompatibility, since self-incompatibility functions in non-distylous Oleaceae species with GA2ox as the sole determinant<ref>{{Cite journal |last=Castric |first=Vincent |last2=Batista |first2=Rita A. |last3=Carré |first3=Amélie |last4=Mousavi |first4=Soraya |last5=Mazoyer |first5=Clément |last6=Godé |first6=Cécile |last7=Gallina |first7=Sophie |last8=Ponitzki |first8=Chloé |last9=Theron |first9=Anthony |last10=Bellec |first10=Arnaud |last11=Marande |first11=William |last12=Santoni |first12=Sylvain |last13=Mariotti |first13=Roberto |last14=Rubini |first14=Andrea |last15=Legrand |first15=Sylvain |date=May 2024 |title=The homomorphic self-incompatibility system in Oleaceae is controlled by a hemizygous genomic region expressing a gibberellin pathway gene |url=https://linkinghub.elsevier.com/retrieve/pii/S0960982224003877 |journal=Current Biology |language=en |volume=34 |issue=9 |pages=1967–1976.e6 |doi=10.1016/j.cub.2024.03.047}}</ref>.

=== The ''S''-locus of ''Fagopyrum'' ===
The S''-''morph of ''Fagopyrum'' contains ~2.8 Mb hemizygous region which likely represents the ''S''-locus as it contains ''S-ELF4'' which establishes female morphology and mating type.<ref name=":14" /><ref name=":17" />

=== The ''S-''locus of ''Gelsemium'' ===
In ''Gelsemium'', the ''S''-locus is composed of four genes, ''GeCYP'', ''GeFRS6'', and ''GeGA3OX'' are hemizygous and ''TAF2'' appears to be allelic with a truncated copy in the L-morph.<ref name=":16" /> ''GeCYP'' appears to share a last common ancestor (or [[ortholog]]) with the ''Primula S''-gene ''CYPT.'' It is currently hypothesized that the for ''S''-genes in ''Gelsemium'' were inherited as a group rather than separately.<ref name=":16" /> This is the only known case of the ''S''-genes being inherited as a group rather than individually.

=== The ''S''-locus of ''Linum'' ===
In ''Linum'' the ''S''-locus is composed of nine genes, two are ''LtTSS1'' and ''LtWDR-44'' the other seven are unnamed and are of unknown function.<ref name=":12" /> ''LtTSS1'' is hypothesized to regulate style length in the S-morph.<ref name=":13" /> [[Synonymous substitution]] analysis of three of the ''S''-genes suggest the ''S''-locus in ''Linum'' evolved in a step by step manner, though only three of the nine genes were analyzed.<ref name=":12" />

=== The ''S''-locus of ''Nymphoides'' ===
The ''S''-locus of ''Nymphoides'' contains three genes ''NinS1'', ''NinKHZ2'', and ''NinBAS1.''<ref name=":18" /> ''NinBAS1'' is only expressed in the style and is hypothetical involved in regulation of brassinosteroids, ''NinS1'' is only expressed in the stamen, ''NinKHZ2'' is expressed in both stamen and style.<ref name=":18" /> Similar to other ''S''-loci, the ''Nymphoides S''-locus appears to have evolved via stepwise duplication events.<ref name=":18" />

=== The ''S''-locus of ''Primula'' ===
In ''Primula'' the ''S-''locus is composed of five genes, ''CYPT''(or ''CYP734A50''), ''GLOT'' (or ''GLOBOSA2'')'', KFBT, PUMT,'' and ''CCMT.'' The supergene evolved in a step-by-step manner, meaning each ''S-''gene duplicated and move to the pre-''S''-locus independently of the others.<ref name=":8">{{Cite journal |last1=Huu |first1=Cuong Nguyen |last2=Keller |first2=Barbara |last3=Conti |first3=Elena |last4=Kappel |first4=Christian |last5=Lenhard |first5=Michael |date=2020-09-15 |title=Supergene evolution via stepwise duplications and neofunctionalization of a floral-organ identity gene |journal=Proceedings of the National Academy of Sciences |language=en |volume=117 |issue=37 |pages=23148–23157 |doi=10.1073/pnas.2006296117 |pmid=32868445 |pmc=7502755 |issn=0027-8424|doi-access=free |bibcode=2020PNAS..11723148H }}</ref><ref name=":9">{{Cite journal |last1=Potente |first1=Giacomo |last2=Léveillé-Bourret |first2=Étienne |last3=Yousefi |first3=Narjes |last4=Choudhury |first4=Rimjhim Roy |last5=Keller |first5=Barbara |last6=Diop |first6=Seydina Issa |last7=Duijsings |first7=Daniël |last8=Pirovano |first8=Walter |last9=Lenhard |first9=Michael |last10=Szövényi |first10=Péter |last11=Conti |first11=Elena |date=2022-02-03 |editor-last=de Meaux |editor-first=Juliette |title=Comparative Genomics Elucidates the Origin of a Supergene Controlling Floral Heteromorphism |url=https://academic.oup.com/mbe/article/doi/10.1093/molbev/msac035/6526404 |journal=Molecular Biology and Evolution |language=en |volume=39 |issue=2 |article-number=msac035 |doi=10.1093/molbev/msac035 |issn=0737-4038 |pmc=8859637 |pmid=35143659}}</ref> Synonymous substitution analysis of the ''S''-genes suggest the oldest ''S''-gene in ''Primula'' is likely ''KFBT'' which likely duplicated about 104 million years ago, followed by ''CYPT''(42.7 MYA),''GLOT'' (37.4 MYA), ''CCMT''(10.3 MYA).<ref name=":9" /> It is unknown when ''PUMT'' evolved as it does not have a [[paralog]] within the ''Primula'' genome.

Of the five ''S''-genes, two have been characterized. ''CYPT,'' a ''[[cytochrome P450]]'' family member, is the female morphology<ref name=":10">{{Cite journal |last1=Huu |first1=Cuong Nguyen |last2=Kappel |first2=Christian |last3=Keller |first3=Barbara |last4=Sicard |first4=Adrien |last5=Takebayashi |first5=Yumiko |last6=Breuninger |first6=Holger |last7=Nowak |first7=Michael D |last8=Bäurle |first8=Isabel |last9=Himmelbach |first9=Axel |last10=Burkart |first10=Michael |last11=Ebbing-Lohaus |first11=Thomas |last12=Sakakibara |first12=Hitoshi |last13=Altschmied |first13=Lothar |last14=Conti |first14=Elena |last15=Lenhard |first15=Michael |date=2016-09-06 |editor-last=Hardtke |editor-first=Christian S |title=Presence versus absence of CYP734A50 underlies the style-length dimorphism in primroses |journal=eLife |volume=5 |article-number=e17956 |doi=10.7554/eLife.17956 |issn=2050-084X |pmc=5012859 |pmid=27596932 |doi-access=free }}</ref> and it is the female self-incompatibility gene,<ref name=":11">{{Cite journal |last1=Huu |first1=Cuong Nguyen |last2=Plaschil |first2=Sylvia |last3=Himmelbach |first3=Axel |last4=Kappel |first4=Christian |last5=Lenhard |first5=Michael |date=2022 |title=Female self-incompatibility type in heterostylous Primula is determined by the brassinosteroid-inactivating cytochrome P450 CYP734A50 |journal=Current Biology |language=en |volume=32 |issue=3 |pages=671–676.e5 |doi=10.1016/j.cub.2021.11.046|pmid=34906354 |s2cid=245128230 |doi-access=free |bibcode=2022CBio...32E.671H }}</ref> meaning it promotes rejection of self pollen. ''CYPT'' is likely producing these phenotypes via inactivation of [[brassinosteroid]]s.<ref name=":10" /><ref name=":11" /> Inactivation of brassinosteroids in the S-morph by ''CYPT'' results in repression of cell elongation in the style by repressing expression of ''PIN5'', ultimately producing the short pistil phenotype.<ref name=":10" /><ref name=":19">{{Cite journal |last1=Liu |first1=Ying |last2=Si |first2=Weijia |last3=Fu |first3=Sitong |last4=Wang |first4=Jia |last5=Cheng |first5=Tangren |last6=Zhang |first6=Qixiang |last7=Pan |first7=Huitang |date=2024-01-08 |title=''PfPIN5'' promotes style elongation by regulating cell length in ''Primula forbesii'' French |journal=Annals of Botany |volume=133 |issue=3 |pages=473–482 |doi=10.1093/aob/mcae004 |pmid=38190350 |pmc=11006536 |issn=0305-7364}}</ref> ''GLOT'' , a ''[[MADS-box|MADS-BOX]]'' family member,<ref>{{Cite journal |last1=Burrows |first1=Benjamin A. |last2=McCubbin |first2=Andrew G. |date=2017 |title=Sequencing the genomic regions flanking S-linked PvGLO sequences confirms the presence of two GLO loci, one of which lies adjacent to the style-length determinant gene CYP734A50 |url=http://link.springer.com/10.1007/s00497-017-0299-9 |journal=Plant Reproduction |language=en |volume=30 |issue=1 |pages=53–67 |doi=10.1007/s00497-017-0299-9 |pmid=28229234 |bibcode=2017PlanR..30...53B |s2cid=22910136 |issn=2194-7953|url-access=subscription }}</ref> ''i''s the male morphology gene as it promotes [[corolla tube]] growth under the stamen.<ref name=":8" /> It is unknown how the other three ''S''-genes are contributing to distyly in ''Primula.''

=== The ''S''-locus of ''Turnera'' ===
In ''Turnera'' the ''S''-locus is composed of three genes, ''BAHD, SPH1,'' and ''YUC6.''<ref name=":4" /> ''BAHD'' is likely an [[acyltransferase]] involved in inactivation of brassinosteroids;<ref name=":15">{{Cite journal |last1=Matzke |first1=Courtney M. |last2=Shore |first2=Joel S. |last3=Neff |first3=Michael M. |last4=McCubbin |first4=Andrew G. |date=2020-11-13 |title=The Turnera Style S-Locus Gene TsBAHD Possesses Brassinosteroid-Inactivating Activity When Expressed in Arabidopsis thaliana |journal=Plants |language=en |volume=9 |issue=11 |page=1566 |doi=10.3390/plants9111566 |issn=2223-7747 |pmc=7697239 |pmid=33202834|doi-access=free |bibcode=2020Plnts...9.1566M }}</ref> it is both the female morphology<ref name=":15" /> and female self-incompatibility gene.<ref>{{Cite journal |last1=Matzke |first1=Courtney M. |last2=Hamam |first2=Hasan J. |last3=Henning |first3=Paige M. |last4=Dougherty |first4=Kyra |last5=Shore |first5=Joel S. |last6=Neff |first6=Michael M. |last7=McCubbin |first7=Andrew G. |date=2021-09-30 |title=Pistil Mating Type and Morphology Are Mediated by the Brassinosteroid Inactivating Activity of the S-Locus Gene BAHD in Heterostylous Turnera Species |journal=International Journal of Molecular Sciences |language=en |volume=22 |issue=19 |article-number=10603 |doi=10.3390/ijms221910603 |issn=1422-0067 |pmc=8509066 |pmid=34638969|doi-access=free }}</ref> ''YUC6'' is likely involved in [[auxin]] biosynthesis based on homology; it is the male self-incompatibility gene and establishes pollen size dimorphisms.<ref>{{Cite journal |last1=Henning |first1=Paige M. |last2=Shore |first2=Joel S. |last3=McCubbin |first3=Andrew G. |date=2022-10-08 |title=The S-Gene YUC6 Pleiotropically Determines Male Mating Type and Pollen Size in Heterostylous Turnera (Passifloraceae): A Novel Neofunctionalization of the YUCCA Gene Family |journal=Plants |language=en |volume=11 |issue=19 |page=2640 |doi=10.3390/plants11192640 |issn=2223-7747 |pmc=9572539 |pmid=36235506|doi-access=free |bibcode=2022Plnts..11.2640H }}</ref> ''SPH1'' is likely involved in filament elongation based on short filament mutant analysis.<ref name=":4" />

== List of families with distylous species ==
Source:<ref name=":3" />

{{columns-list|colwidth=25em|
* [[Acanthaceae]]
* [[Amaryllidaceae]]
* [[Boraginaceae]]
* [[Connaraceae]]
* [[Erythroxylaceae]]
* [[Fabaceae]]
* [[Gelsemiaceae]]
* [[Gentianaceae]]
* [[Hypericaceae]]
* [[Iridaceae]]
* [[Lamiaceae]]
* [[Linaceae]]
* [[Lythraceae]]
* [[Malvaceae]]
* [[Menyanthaceae]]
* [[Oleaceae]]
* [[Oxalidaceae]]
* [[Passifloraceae]]
* [[Plumbaginaceae]]
* [[Polemoniaceae]]
* [[Polygonaceae]]
* [[Pontederiaceae]]
* [[Primulaceae]]
* [[Rubiaceae]]
* [[Santalaceae]]
* [[Saxifragaceae]]
* [[Schoepfiaceae]]
* [[Thymelaeaceae]]
}}

== References ==
<references />

[[Category:Plant reproduction]]
[[Category:Plant morphology]]
[[Category:Pollination]]
[[Category:Genetics]]
[[Category:Evolution]]

Distyly - История изменений

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