Sex-determination system

{{Expand section|date=June 2021}}

Sex determination was discovered in the mealworm by the American geneticist Nettie Stevens in 1903.{{Cite web|url=https://www.nature.com/scitable/topicpage/nettie-stevens-a-discoverer-of-sex-chromosomes-6580266|title=Nettie Stevens: A Discoverer of Sex Chromosomes {{!}} Learn Science at Scitable|website=www.nature.com|access-date=2018-06-07|archive-date=7 April 2019|archive-url=https://web.archive.org/web/20190407152710/https://www.nature.com/scitable/topicpage/nettie-stevens-a-discoverer-of-sex-chromosomes-6580266|url-status=live}}{{cite journal | vauthors = Ogilvie MB, Choquette CJ | title = Nettie Maria Stevens (1861–1912): her life and contributions to cytogenetics | journal = Proceedings of the American Philosophical Society | volume = 125 | issue = 4 | pages = 292–311 | date = August 1981 | pmid = 11620765 | jstor = 986332 }}{{Cite web|url=https://embryo.asu.edu/pages/nettie-maria-stevens-1861-1912|title=Nettie Maria Stevens (1861–1912) {{!}} The Embryo Project Encyclopedia|website=embryo.asu.edu|access-date=2018-06-07|archive-date=8 April 2019|archive-url=https://web.archive.org/web/20190408010235/https://embryo.asu.edu/pages/nettie-maria-stevens-1861-1912|url-status=live}}

In 1694, J.R. Camerarius, conducted early experiments on pollination and reported the existence of male and female characteristics in plants(Maize).

In 1866, Gregor Mendel published on inheritance of genetic traits. This is known as Mendelian inheritance and it eventually established the modern understanding of inheritance from two gametes.

In 1902, C.E. McClung identified sex chromosomes in bugs.

In 1917, C.E. Allen, discovered sex determination mechanisms in plants.

In 1922, C.B. Bridges, put forth the Genic Balance Theory of sex determination.

In 1928, Fritz Baltzer first described environmental sex determination.{{Cite journal |last=Baltzer |first=Fritz |date=1928 |title=Über metagame Geschlechtsbestimmung und ihre Beziehung zu einigen Problemen der Entwicklungsmechanik und Vererbung (Auf Grund von Versuchen an Bonellia) |journal=Verhandlungen der Deutschen Zoologischen Gesellschaft 32: 273-325 |language=de |volume=32 |pages=273-325}}

Among animals, the most common chromosomal sex determination systems are XY, XO, ZW, ZO, but with numerous exceptions.

According to the Tree of Sex database{{Cite journal |date=2014-06-24 |title=Tree of Sex: A database of sexual systems |journal=Scientific Data |volume=1 |pages=140015 |doi=10.1038/sdata.2014.15 |issn=2052-4463 |pmc=4322564 |pmid=25977773 |last1=Ashman |first1=Tia-Lynn |last2=Bachtrog |first2=Doris |last3=Blackmon |first3=Heath |last4=Goldberg |first4=Emma E. |last5=Hahn |first5=Matthew W. |last6=Kirkpatrick |first6=Mark |last7=Kitano |first7=Jun |last8=Mank |first8=Judith E. |last9=Mayrose |first9=Itay |last10=Ming |first10=Ray |last11=Otto |first11=Sarah P. |last12=Peichel |first12=Catherine L. |last13=Pennell |first13=Matthew W. |last14=Perrin |first14=Nicolas |last15=Ross |first15=Laura |last16=Valenzuela |first16=Nicole |last17=Vamosi |first17=Jana C. }} (as of 2023), the known sex determination systems are:{{Cite journal |last1=Hayashi |first1=Shun |last2=Abe |first2=Takuya |last3=Igawa |first3=Takeshi |last4=Katsura |first4=Yukako |last5=Kazama |first5=Yusuke |last6=Nozawa |first6=Masafumi |date=2024-07-31 |title=Sex chromosome cycle as a mechanism of stable sex determination |url=https://academic.oup.com/jb/article/176/2/81/7709515 |journal=The Journal of Biochemistry |volume=176 |issue=2 |pages=81–95 |doi=10.1093/jb/mvae045 |issn=0021-924X |pmc=11289310 |pmid=38982631}}

^{1. complex sex chromosomes, homomorphic sex chromosomes, or others}

File:Drosophila XY sex-determination.svg

File:Human male karyotpe high resolution - XY chromosome cropped.JPG]]

{{main|XY sex-determination system}}

The XX/XY sex-determination system is the most familiar, as it is found in humans. The XX/XY system is found in most other mammals, as well as some insects. In this system, females have two of the same kind of sex chromosome (XX), while males have two distinct sex chromosomes (XY). The X and Y sex chromosomes are different in shape and size from each other, unlike the rest of the chromosomes (autosomes), and are sometimes called allosomes. In some species, such as humans, organisms remain sex indifferent for a time during development (embryogenesis); in others, however, such as fruit flies, sexual differentiation occurs as soon as the egg is fertilized.

Some species (including humans) have a gene SRY on the Y chromosome that determines maleness. Members of SRY-reliant species can have uncommon XY chromosomal combinations such as XXY and still live.{{cite journal |vauthors=Hake L |title=Genetic Mechanisms of Sex Determination |journal=Nature Education |year=2008 |volume=1 |issue=1 |url=http://www.nature.com/scitable/topicpage/genetic-mechanisms-of-sex-determination-314 |access-date=8 December 2011 |archive-date=19 August 2017 |archive-url=https://web.archive.org/web/20170819121941/http://www.nature.com/scitable/topicpage/genetic-mechanisms-of-sex-determination-314 |url-status=live }}

Human sex is determined by the presence or absence of a Y chromosome with a functional SRY gene. Once the SRY gene is activated, cells create testosterone and anti-müllerian hormone which typically ensures the development of a single, male reproductive system. In typical XX embryos, cells secrete estrogen, which drives the body toward the female pathway.

In Y-centered sex determination, the SRY gene is the main gene in determining male characteristics, but multiple genes are required to develop testes. In XY mice, lack of the gene DAX1 on the X chromosome results in sterility, but in humans it causes adrenal hypoplasia congenita.{{cite journal | vauthors = Goodfellow PN, Camerino G | title = DAX-1, an 'antitestis' gene | journal = Cellular and Molecular Life Sciences | volume = 55 | issue = 6–7 | pages = 857–863 | date = June 1999 | pmid = 10412368 | doi = 10.1007/PL00013201 | s2cid = 19764423 | pmc = 11147076 }} However, when an extra DAX1 gene is placed on the X chromosome, the result is a female, despite the existence of SRY, since it overrides the effects of SRY.{{cite journal |author=Chandra, H. S. |title=Another way of looking at the enigma of sex determination in Ellobius lutescens |journal=Current Science |date=25 April 1999 |page=1072 |volume=76 |issue=8}} Even when there are normal sex chromosomes in XX females, duplication or expression of SOX9 causes testes to develop.{{cite journal | vauthors = Cox JJ, Willatt L, Homfray T, Woods CG | title = A SOX9 duplication and familial 46,XX developmental testicular disorder | journal = The New England Journal of Medicine | volume = 364 | issue = 1 | pages = 91–93 | date = January 2011 | pmid = 21208124 | doi = 10.1056/NEJMc1010311 | doi-access = free }}{{cite journal | vauthors = Huang B, Wang S, Ning Y, Lamb AN, Bartley J | title = Autosomal XX sex reversal caused by duplication of SOX9 | journal = American Journal of Medical Genetics | volume = 87 | issue = 4 | pages = 349–353 | date = December 1999 | pmid = 10588843 | doi = 10.1002/(SICI)1096-8628(19991203)87:4<349::AID-AJMG13>3.0.CO;2-N }} Gradual sex reversal in developed mice can also occur when the gene FOXL2 is removed from females.{{cite journal | vauthors = Uhlenhaut NH, Jakob S, Anlag K, Eisenberger T, Sekido R, Kress J, Treier AC, Klugmann C, Klasen C, Holter NI, Riethmacher D, Schütz G, Cooney AJ, Lovell-Badge R, Treier M | display-authors = 6 | title = Somatic sex reprogramming of adult ovaries to testes by FOXL2 ablation | journal = Cell | volume = 139 | issue = 6 | pages = 1130–1142 | date = December 2009 | pmid = 20005806 | doi = 10.1016/j.cell.2009.11.021 | doi-access = free }} Even though the gene DMRT1 is used by birds as their sex locus, species who have XY chromosomes also rely upon DMRT1, contained on chromosome 9, for sexual differentiation at some point in their formation.

Some species, such as fruit flies, use the presence of two X chromosomes to determine femaleness.{{cite journal | vauthors = Penalva LO, Sánchez L | title = RNA binding protein sex-lethal (Sxl) and control of Drosophila sex determination and dosage compensation | journal = Microbiology and Molecular Biology Reviews | volume = 67 | issue = 3 | pages = 343–59, table of contents | date = September 2003 | pmid = 12966139 | pmc = 193869 | doi = 10.1128/MMBR.67.3.343-359.2003 }} Species that use the number of Xs to determine sex are nonviable with an extra X chromosome.

Some fish have variants of the XY sex-determination system, as well as the regular system. For example, while having an XY format, Xiphophorus nezahualcoyotl and X. milleri also have a second Y chromosome, known as Y', that creates XY' females and YY' males.{{cite journal | vauthors = Schartl M | title = A comparative view on sex determination in medaka | journal = Mechanisms of Development | volume = 121 | issue = 7–8 | pages = 639–645 | date = July 2004 | pmid = 15210173 | doi = 10.1016/j.mod.2004.03.001 | s2cid = 17401686 | doi-access = free }}

At least one monotreme, the platypus, presents a particular sex determination scheme that in some ways resembles that of the ZW sex chromosomes of birds and lacks the SRY gene. The platypus has sex chromosomes $X\_1,\; X\_2,\; X\_3,\; X\_4,\; X\_5,\; Y\_1,\; Y\_2,\; Y\_3,\; Y\_4,\; Y\_5$. The males have $X\_1Y\_1/X\_2Y\_2/X\_3Y\_3/X\_4Y\_4/X\_5Y\_5$, while females have $X\_1X\_1/X\_2X\_2/X\_3X\_3/X\_4X\_4/X\_5X\_5$. During meiosis, 5 of X form one chain, and 5 of Y form another chain. Thus, they behave effectively as a typical XY chromosomal system, except each of X and Y is broken into 5 parts, with the effect that recombinations occur very frequently at 4 particular points.{{Cite journal |last1=Gruetzner |first1=Frank |last2=Ashley |first2=Terry |last3=Rowell |first3=David M. |last4=Marshall Graves |first4=Jennifer A. |date=2006-04-01 |title=How did the platypus get its sex chromosome chain? A comparison of meiotic multiples and sex chromosomes in plants and animals |url=https://doi.org/10.1007/s00412-005-0034-4 |journal=Chromosoma |volume=115 |issue=2 |pages=75–88 |doi=10.1007/s00412-005-0034-4 |pmid=16344965 |issn=1432-0886}} One of the X chromosomes is homologous to the human X chromosome, and another is homologous to the bird Z chromosome.{{Cite journal |last1=Grützner |first1=Frank |last2=Rens |first2=Willem |last3=Tsend-Ayush |first3=Enkhjargal |last4=El-Mogharbel |first4=Nisrine |last5=O'Brien |first5=Patricia C. M. |last6=Jones |first6=Russell C. |last7=Ferguson-Smith |first7=Malcolm A. |last8=Marshall Graves |first8=Jennifer A. |date=December 2004 |title=In the platypus a meiotic chain of ten sex chromosomes shares genes with the bird Z and mammal X chromosomes |url=https://www.nature.com/articles/nature03021 |journal=Nature |volume=432 |issue=7019 |pages=913–917 |doi=10.1038/nature03021 |pmid=15502814 |bibcode=2004Natur.432..913G |issn=1476-4687}}

Although it is an XY system, the platypus' sex chromosomes share no homologues with eutherian sex chromosomes.{{cite journal | vauthors = Warren WC, Hillier LW, Marshall Graves JA, Birney E, Ponting CP, Grützner F, Belov K, Miller W, Clarke L, Chinwalla AT, Yang SP, Heger A, Locke DP, Miethke P, Waters PD, Veyrunes F, Fulton L, Fulton B, Graves T, Wallis J, Puente XS, López-Otín C, Ordóñez GR, Eichler EE, Chen L, Cheng Z, Deakin JE, Alsop A, Thompson K, Kirby P, Papenfuss AT, Wakefield MJ, Olender T, Lancet D, Huttley GA, Smit AF, Pask A, Temple-Smith P, Batzer MA, Walker JA, Konkel MK, Harris RS, Whittington CM, Wong ES, Gemmell NJ, Buschiazzo E, Vargas Jentzsch IM, Merkel A, Schmitz J, Zemann A, Churakov G, Kriegs JO, Brosius J, Murchison EP, Sachidanandam R, Smith C, Hannon GJ, Tsend-Ayush E, McMillan D, Attenborough R, Rens W, Ferguson-Smith M, Lefèvre CM, Sharp JA, Nicholas KR, Ray DA, Kube M, Reinhardt R, Pringle TH, Taylor J, Jones RC, Nixon B, Dacheux JL, Niwa H, Sekita Y, Huang X, Stark A, Kheradpour P, Kellis M, Flicek P, Chen Y, Webber C, Hardison R, Nelson J, Hallsworth-Pepin K, Delehaunty K, Markovic C, Minx P, Feng Y, Kremitzki C, Mitreva M, Glasscock J, Wylie T, Wohldmann P, Thiru P, Nhan MN, Pohl CS, Smith SM, Hou S, Nefedov M, de Jong PJ, Renfree MB, Mardis ER, Wilson RK | display-authors = 6 | title = Genome analysis of the platypus reveals unique signatures of evolution | journal = Nature | volume = 453 | issue = 7192 | pages = 175–183 | date = May 2008 | pmid = 18464734 | pmc = 2803040 | doi = 10.1038/nature06936 | bibcode = 2008Natur.453..175W }} Instead, homologues with eutherian sex chromosomes lie on the platypus chromosome 6, which means that the eutherian sex chromosomes were autosomes at the time that the monotremes diverged from the therian mammals (marsupials and eutherian mammals). However, homologues to the avian DMRT1 gene on platypus sex chromosomes X3 and X5 suggest that it is possible the sex-determining gene for the platypus is the same one that is involved in bird sex-determination. More research must be conducted in order to determine the exact sex determining gene of the platypus.{{cite journal | vauthors = Gruetzner F, Ashley T, Rowell DM, Marshall Graves JA | title = How did the platypus get its sex chromosome chain? A comparison of meiotic multiples and sex chromosomes in plants and animals | journal = Chromosoma | volume = 115 | issue = 2 | pages = 75–88 | date = April 2006 | pmid = 16344965 | doi = 10.1007/s00412-005-0034-4 | name-list-style = amp | s2cid = 23603889 }}

File:Critique of the Theory of Evolution Fig 060.svg

{{Main |X0 sex-determination system}}

In this variant of the XY system, females have two copies of the sex chromosome (XX) but males have only one (X0). The 0 denotes the absence of a second sex chromosome. Generally in this method, the sex is determined by amount of genes expressed across the two chromosomes. This system is observed in a number of insects, including the grasshoppers and crickets of order Orthoptera and in cockroaches (order Blattodea). A small number of mammals also lack a Y chromosome. These include the Amami spiny rat (Tokudaia osimensis) and the Tokunoshima spiny rat (Tokudaia tokunoshimensis) and Sorex araneus, a shrew species. Transcaucasian mole voles (Ellobius lutescens) also have a form of XO determination, in which both sexes lack a second sex chromosome. The mechanism of sex determination is not yet understood.{{cite journal | vauthors = Kuroiwa A, Handa S, Nishiyama C, Chiba E, Yamada F, Abe S, Matsuda Y | title = Additional copies of CBX2 in the genomes of males of mammals lacking SRY, the Amami spiny rat (Tokudaia osimensis) and the Tokunoshima spiny rat (Tokudaia tokunoshimensis) | journal = Chromosome Research | volume = 19 | issue = 5 | pages = 635–644 | date = July 2011 | pmid = 21656076 | doi = 10.1007/s10577-011-9223-6 | s2cid = 23311263 }}

The nematode C. elegans is male with one sex chromosome (X0); with a pair of chromosomes (XX) it is a hermaphrodite.{{Cite book | vauthors = Majerus ME | title = Sex wars: genes, bacteria, and biased sex ratios | publisher = Princeton University Press | pages = 250 | url = https://books.google.com/books?id=vDHOYPQ2mmYC&q=zo,+zww,+zzww+lepidoptera | isbn = 978-0-691-00981-0 | access-date = 4 November 2011 | year = 2003 }}

Its main sex gene is XOL, which encodes XOL-1 and also controls the expression of the genes TRA-2 and HER-1. These genes reduce male gene activation and increase it, respectively.{{cite journal | vauthors = Kuwabara PE, Okkema PG, Kimble J | title = tra-2 encodes a membrane protein and may mediate cell communication in the Caenorhabditis elegans sex determination pathway | journal = Molecular Biology of the Cell | volume = 3 | issue = 4 | pages = 461–473 | date = April 1992 | pmid = 1498366 | pmc = 275596 | doi = 10.1091/mbc.3.4.461 }}

{{Main |ZW sex-determination system}}

The ZW sex-determination system is found in birds, some reptiles, and some insects and other organisms. The ZW sex-determination system is reversed compared to the XY system: females have two different kinds of chromosomes (ZW), and males have two of the same kind of chromosomes (ZZ). In the chicken, this was found to be dependent on the expression of DMRT1.{{cite journal | vauthors = Smith CA, Roeszler KN, Ohnesorg T, Cummins DM, Farlie PG, Doran TJ, Sinclair AH | title = The avian Z-linked gene DMRT1 is required for male sex determination in the chicken | journal = Nature | volume = 461 | issue = 7261 | pages = 267–271 | date = September 2009 | pmid = 19710650 | doi = 10.1038/nature08298 | s2cid = 4413389 | bibcode = 2009Natur.461..267S }} In birds, the genes FET1 and ASW are found on the W chromosome for females, similar to how the Y chromosome contains SRY. However, not all species depend upon the W for their sex. For example, there are moths and butterflies that are ZW, but some have been found female with ZO, as well as female with ZZW. Also, while mammals deactivate one of their extra X chromosomes when female, it appears that in the case of Lepidoptera, the males produce double the normal amount of enzymes, due to having two Z's. Because the use of ZW sex determination is varied, it is still unknown how exactly most species determine their sex. However, reportedly, the silkworm Bombyx mori uses a single female-specific piRNA as the primary determiner of sex.{{cite journal | vauthors = Kiuchi T, Koga H, Kawamoto M, Shoji K, Sakai H, Arai Y, Ishihara G, Kawaoka S, Sugano S, Shimada T, Suzuki Y, Suzuki MG, Katsuma S | display-authors = 6 | title = A single female-specific piRNA is the primary determiner of sex in the silkworm | journal = Nature | volume = 509 | issue = 7502 | pages = 633–636 | date = May 2014 | pmid = 24828047 | doi = 10.1038/nature13315 | s2cid = 205238635 | bibcode = 2014Natur.509..633K }} Despite the similarities between the ZW and XY systems, these sex chromosomes evolved separately. In the case of the chicken, their Z chromosome is more similar to humans' autosome 9.{{cite journal | vauthors = Stiglec R, Ezaz T, Graves JA | title = A new look at the evolution of avian sex chromosomes | journal = Cytogenetic and Genome Research | volume = 117 | issue = 1–4 | pages = 103–109 | year = 2007 | pmid = 17675850 | doi = 10.1159/000103170 | s2cid = 12932564 }} The chicken's Z chromosome also seems to be related to the X chromosome of the platypus.{{cite journal | vauthors = Grützner F, Rens W, Tsend-Ayush E, El-Mogharbel N, O'Brien PC, Jones RC, Ferguson-Smith MA, Marshall Graves JA | display-authors = 6 | title = In the platypus a meiotic chain of ten sex chromosomes shares genes with the bird Z and mammal X chromosomes | journal = Nature | volume = 432 | issue = 7019 | pages = 913–917 | date = December 2004 | pmid = 15502814 | doi = 10.1038/nature03021 | name-list-style = amp | s2cid = 4379897 | bibcode = 2004Natur.432..913G }} When a ZW species, such as the Komodo dragon, reproduces parthenogenetically, usually only males are produced. This is due to the fact that the haploid eggs double their chromosomes, resulting in ZZ or WW. The ZZ become males, but the WW are not viable and are not brought to term.{{cite news |work=BBC News |url=http://news.bbc.co.uk/2/hi/science/nature/6196225.stm |title=Virgin births for giant lizards |access-date=13 March 2008 |date=20 December 2006 |archive-date=4 November 2014 |archive-url=https://web.archive.org/web/20141104002358/http://news.bbc.co.uk/2/hi/science/nature/6196225.stm |url-status=live }}

In both XY and ZW sex determination systems, the sex chromosome carrying the critical factors is often significantly smaller, carrying little more than the genes necessary for triggering the development of a given sex.{{cite web|title=Evolution of the Y Chromosome|url=http://www.learner.org/channel/courses/biology/textbook/gender/gender_4.html|url-status=dead|archive-url=https://web.archive.org/web/20041104181945/http://www.learner.org/channel/courses/biology/textbook/gender/gender_4.html|archive-date=November 4, 2004|access-date=1 April 2008|website=Annenberg Media}}{{Better source needed|date=May 2021}}

{{Main |Z0 sex-determination system}}

The ZZ/Z0 sex-determination system is found in some moths. In these insects there is one sex chromosome, Z. Males have two Z chromosomes, whereas females have one Z. Males are ZZ, while females are Z0.{{cite journal | vauthors = Traut W, Sahara K, Marec F | title = Sex chromosomes and sex determination in Lepidoptera | journal = Sexual Development | volume = 1 | issue = 6 | pages = 332–346 | date = 2007 | pmid = 18391545 | doi = 10.1159/000111765 | s2cid = 6885122 }}{{cite web|url=http://www.nature.com/scitable/topicpage/genetic-mechanisms-of-sex-determination-314|title=Genetic Mechanisms of Sex Determination - Learn Science at Scitable|website=www.nature.com|access-date=8 December 2011|archive-date=19 August 2017|archive-url=https://web.archive.org/web/20170819121941/http://www.nature.com/scitable/topicpage/genetic-mechanisms-of-sex-determination-314|url-status=live}}{{cite book|url=https://books.google.com/books?id=5w8FgSGuH34C&q=ZO+sex-determination+system+moth&pg=PA461|title=Handbuch Der Zoologie / Handbook of Zoology|publisher=Walter de Gruyter|via=Google Books|isbn=9783110162103|year=1925}}

In some bryophyte and some algae species, the gametophyte stage of the life cycle, rather than being hermaphrodite, occurs as separate male or female individuals that produce male and female gametes respectively. When meiosis occurs in the sporophyte generation of the life cycle, the sex chromosomes known as U and V assort in spores that carry either the U chromosome and give rise to female gametophytes, or the V chromosome and give rise to male gametophytes.{{cite journal | vauthors = Bachtrog D, Kirkpatrick M, Mank JE, McDaniel SF, Pires JC, Rice W, Valenzuela N | title = Are all sex chromosomes created equal? | journal = Trends in Genetics | volume = 27 | issue = 9 | pages = 350–357 | date = September 2011 | pmid = 21962970 | doi = 10.1016/j.tig.2011.05.005 }}{{cite journal |author1=Renner, S. S. |author2=Heinrichs, J. |author3=Sousa, A. |year=2017 |title= The sex chromosomes of bryophytes: Recent insights, open questions, and reinvestigations of Frullania dilatata and Plagiochila asplenioides. |journal=Journal of Systematics and Evolution |volume=55 |issue=4 |pages=333–339 |doi=10.1111/jse.12266|doi-access=free }}

{{Main|Mating type}}

The mating type in microorganisms is analogous to sex in multi-cellular organisms, and is sometimes described using those terms, though they are not necessarily correlated with physical body structures. Some species have more than two mating types. Tetrahymena, a type of ciliate, has 7 mating types

Mating types are extensively studied in fungi. Among fungi, mating type is determined by chromosomal regions called mating-type loci. Furthermore, it is not as simple as "two different mating types can mate", but rather, a matter of combinatorics. As a simple example, most basidiomycete have a "tetrapolar heterothallism" mating system: there are two loci, and mating between two individuals is possible if the alleles on both loci are different. For example, if there are 3 alleles per locus, then there would be 9 mating types, each of which can mate with 4 other mating types.{{Cite journal |last1=Idnurm |first1=Alexander |last2=Hood |first2=Michael E. |last3=Johannesson |first3=Hanna |last4=Giraud |first4=Tatiana |date=2015-12-01 |title=Contrasted patterns in mating-type chromosomes in fungi: Hotspots versus coldspots of recombination |journal=Fungal Biology Reviews |series=Special Issue: Fungal sex and mushrooms – A credit to Lorna Casselton |volume=29 |issue=3 |pages=220–229 |doi=10.1016/j.fbr.2015.06.001 |pmid=26688691 |bibcode=2015FunBR..29..220I |issn=1749-4613|pmc=4680991 }} By multiplicative combination, it generates a vast number of mating types. For example, Schizophyllum commune, a type of fungus, has $9\; \backslash times\; 32\; \backslash times\; 9\; \backslash times\; 9\; =\; 23328$ mating types.File:Haplodiploid-sex-determination-system3.png

{{main|Haplodiploidy}}

Haplodiploidy is found in insects belonging to Hymenoptera, such as ants and bees. Sex determination is controlled by the zygosity of a complementary sex determiner (csd) locus. Unfertilized eggs develop into haploid individuals which have a single, hemizygous copy of the csd locus and are therefore males. Fertilized eggs develop into diploid individuals which, due to high variability in the csd locus, are generally heterozygous females. In rare instances diploid individuals may be homozygous, these develop into sterile males.

The gene acting as a csd locus has been identified in the honeybee and several candidate genes have been proposed as a csd locus for other Hymenopterans.{{cite journal | vauthors = Beye M, Hasselmann M, Fondrk MK, Page RE, Omholt SW | title = The gene csd is the primary signal for sexual development in the honeybee and encodes an SR-type protein | journal = Cell | volume = 114 | issue = 4 | pages = 419–429 | date = August 2003 | pmid = 12941271 | doi = 10.1016/S0092-8674(03)00606-8 | doi-access = free }}{{cite journal | vauthors = Privman E, Wurm Y, Keller L | title = Duplication and concerted evolution in a master sex determiner under balancing selection | journal = Proceedings. Biological Sciences | volume = 280 | issue = 1758 | pages = 20122968 | date = May 2013 | pmid = 23466984 | pmc = 3619454 | doi = 10.1098/rspb.2012.2968 | doi-access = free }}{{cite journal | vauthors = Miyakawa MO, Tsuchida K, Miyakawa H | title = The doublesex gene integrates multi-locus complementary sex determination signals in the Japanese ant, Vollenhovia emeryi | journal = Insect Biochemistry and Molecular Biology | volume = 94 | pages = 42–49 | date = March 2018 | pmid = 29408414 | doi = 10.1016/j.ibmb.2018.01.006 | bibcode = 2018IBMB...94...42M }}

Most females in the Hymenoptera order can decide the sex of their offspring by holding received sperm in their spermatheca and either releasing it into their oviduct or not. This allows them to create more workers, depending on the status of the colony.{{cite journal | vauthors = van Wilgenburg E, Driessen G, Beukeboom LW | title = Single locus complementary sex determination in Hymenoptera: an "unintelligent" design? | journal = Frontiers in Zoology | volume = 3 | issue = 1 | pages = 1 | date = January 2006 | pmid = 16393347 | pmc = 1360072 | doi = 10.1186/1742-9994-3-1 | doi-access = free }}

Polygenic sex determination is when the sex is primarily determined by genes that occur on multiple non-homologous chromosomes. The environment may have a limited, minor influence on sex determination. Examples include African cichlid fish (Metriaclima spp.), lemmings (Myopus schisticolor), green swordtail, medaka, etc. In such systems, there is typically a dominance hierarchy, where one system is dominant over another if in conflict. For example, in some species of cichlid fish from Lake Malawi, if an individual has both the XY locus (on one chromosome pair) and the WZ locus (on another chromosome pair), then the W is dominant and the individual has a female phenotype.{{Cite journal |last1=Moore |first1=Emily C. |last2=Roberts |first2=Reade B. |date=June 2013 |title=Polygenic sex determination |url=https://linkinghub.elsevier.com/retrieve/pii/S0960982213004120 |journal=Current Biology |volume=23 |issue=12 |pages=R510–R512 |doi=10.1016/j.cub.2013.04.004|pmid=23787041 |bibcode=2013CBio...23.R510M }}

The sex-determination system of zebrafish is polygenic. Juvenile zebrafishes (0–30 days after hatching) have both ovary-like tissue to testis tissue. They then develop into male or female adults, with the determination based on a complex interaction genes on multiple chromosomes, but not affected by environmental variations.{{Cite journal |last1=Liew |first1=Woei Chang |last2=Bartfai |first2=Richard |last3=Lim |first3=Zijie |last4=Sreenivasan |first4=Rajini |last5=Siegfried |first5=Kellee R. |last6=Orban |first6=Laszlo |date=2012-04-10 |editor-last=Alsina |editor-first=Berta |title=Polygenic Sex Determination System in Zebrafish |journal=PLOS ONE |volume=7 |issue=4 |pages=e34397 |doi=10.1371/journal.pone.0034397 |doi-access=free |issn=1932-6203 |pmc=3323597 |pmid=22506019|bibcode=2012PLoSO...734397L }}{{Cite journal |last1=Liew |first1=Woei Chang |last2=Orbán |first2=László |date=March 2014 |title=Zebrafish sex: a complicated affair |journal=Briefings in Functional Genomics |volume=13 |issue=2 |pages=172–187 |doi=10.1093/bfgp/elt041 |issn=2041-2649 |pmc=3954038 |pmid=24148942}}

In systems with two sex chromosomes, they can be heteromorphic or homomorphic. Homomorphic sex chromosomes are almost identical in size and gene content. The two familiar kinds of sex chromosome pairs (XY and ZW) are heteromorphic. Homomorphic sex chromosomes exist among pufferfish, ratite birds, pythons, and European tree frogs. Some are quite old, meaning that there is some evolutionary force that resists their differentiation.{{Cite journal |last1=Wright |first1=Alison E. |last2=Dean |first2=Rebecca |last3=Zimmer |first3=Fabian |last4=Mank |first4=Judith E. |date=2016-07-04 |title=How to make a sex chromosome |journal=Nature Communications |volume=7 |issue=1 |pages=12087 |doi=10.1038/ncomms12087 |pmid=27373494 |pmc=4932193 |bibcode=2016NatCo...712087W |issn=2041-1723}} For example, three species of European tree frogs have homologous, homomorphic sex chromosomes, and this homomorphism was maintained for at least 5.4 million years by occasional recombination.{{Cite journal |last1=Stöck |first1=Matthias |last2=Horn |first2=Agnès |last3=Grossen |first3=Christine |last4=Lindtke |first4=Dorothea |last5=Sermier |first5=Roberto |last6=Betto-Colliard |first6=Caroline |last7=Dufresnes |first7=Christophe |last8=Bonjour |first8=Emmanuel |last9=Dumas |first9=Zoé |last10=Luquet |first10=Emilien |last11=Maddalena |first11=Tiziano |last12=Sousa |first12=Helena Clavero |last13=Martinez-Solano |first13=Iñigo |last14=Perrin |first14=Nicolas |date=2011-05-17 |editor-last=Rice |editor-first=William R. |title=Ever-Young Sex Chromosomes in European Tree Frogs |journal=PLOS Biology |volume=9 |issue=5 |pages=e1001062 |doi=10.1371/journal.pbio.1001062 |doi-access=free |issn=1545-7885 |pmc=3100596 |pmid=21629756}}

The Nematocera, particularly the Simuliids and Chironomus, have sex determination regions that are labile, meaning that one species may have the sex determination region in one chromosome, but a closely related species might have the same region moved to a different non-homologous chromosome. Some species even have the sex determination region different among individuals within the same species (intraspecific variation).{{Cite journal |last1=Furman |first1=Benjamin L S |last2=Metzger |first2=David C H |last3=Darolti |first3=Iulia |last4=Wright |first4=Alison E |last5=Sandkam |first5=Benjamin A |last6=Almeida |first6=Pedro |last7=Shu |first7=Jacelyn J |last8=Mank |first8=Judith E |date=2020-06-01 |editor-last=Fraser |editor-first=Bonnie |title=Sex Chromosome Evolution: So Many Exceptions to the Rules |url=https://academic.oup.com/gbe/article/12/6/750/5823304 |journal=Genome Biology and Evolution |volume=12 |issue=6 |pages=750–763 |doi=10.1093/gbe/evaa081 |issn=1759-6653 |pmc=7268786 |pmid=32315410}}{{Cite journal |last1=Martin |first1=Jon |last2=Lee |first2=B. T. O. |date=September 1984 |title=A phylogenetic study of sex determiner location in a group of Australasian Chironomus species (Diptera, Chironomidae) |url=http://link.springer.com/10.1007/BF00292396 |journal=Chromosoma |volume=90 |issue=3 |pages=190–197 |doi=10.1007/BF00292396 |issn=0009-5915}}{{Cite journal |last1=Martin |first1=Jon |last2=Kuvangkadilok |first2=Chaliow |last3=Peart |first3=Dianne H. |last4=Lee |first4=Barry T. O. |date=June 1980 |title=Multiple sex determining regions in a group of related Chironomus species (Diptera:Chironomidae) |url=https://www.nature.com/articles/hdy198034 |journal=Heredity |volume=44 |issue=3 |pages=367–382 |doi=10.1038/hdy.1980.34 |issn=1365-2540}} In some species, some populations have homomorphic sex chromosomes while other populations have heteromorphic sex chromosomes.

The New Zealand frog, Leiopelma hochstetteri, uses a supernumerary sex chromosome. With zero of that chromosome, the frog develops into a male. With one or more, the frog develops into a female. One female had as many as 16 of that chromosome.{{Cite journal |last=Green |first=David M. |date=1988-09-01 |title=Cytogenetics of the endemic New Zealand frog, Leiopelma hochstetteri: extraordinary supernumerary chromosome variation and a unique sex-chromosome system |url=https://doi.org/10.1007/BF00331795 |journal=Chromosoma |volume=97 |issue=1 |pages=55–70 |doi=10.1007/BF00331795 |issn=1432-0886}}

Different populations of the Japanese frog Rana rugosa uses different systems. Two use homomorphic male heterogamety, one uses XX/XY, one uses ZZ/ZW. Remarkably, the X and Z chromosomes are homologous, and the Y and W as well. Dmrt1 is on autosome 1 and not sex-linked. This means that an XX female individual is genetically similar to a ZZ male individual, and an XY male individual is to a ZW female individual. The mechanism behind this is yet unclear, but it is hypothesized that during its recent evolution, the XY-to-ZW transition occurred twice.{{Cite journal |last1=Uno |first1=Yoshinobu |last2=Nishida |first2=Chizuko |last3=Oshima |first3=Yuki |last4=Yokoyama |first4=Satoshi |last5=Miura |first5=Ikuo |last6=Matsuda |first6=Yoichi |last7=Nakamura |first7=Masahisa |date=June 2008 |title=Comparative chromosome mapping of sex-linked genes and identification of sex chromosomal rearrangements in the Japanese wrinkled frog (Rana rugosa, Ranidae) with ZW and XY sex chromosome systems |url=https://link.springer.com/10.1007/s10577-008-1217-7 |journal=Chromosome Research |volume=16 |issue=4 |pages=637–647 |doi=10.1007/s10577-008-1217-7 |pmid=18484182 |issn=0967-3849}}{{Cite journal |last=Graves |first=Jennifer A. Marshall |date=2008-12-01 |title=Weird Animal Genomes and the Evolution of Vertebrate Sex and Sex Chromosomes |url=http://dx.doi.org/10.1146/annurev.genet.42.110807.091714 |journal=Annual Review of Genetics |volume=42 |issue=1 |pages=565–586 |doi=10.1146/annurev.genet.42.110807.091714 |issn=0066-4197}}

Clarias gariepinus uses both XX/XY and ZW/ZZ system within the species, with some populations using homomorphic XX/XY while others using heteromorphic ZW/ZZ. A population in Thailand appears to use both systems simultaneously, possibly because C. gariepinus were not native to Thailand, and were introduced from different source populations which resulted in a mixture.{{Cite journal |last1=Nguyen |first1=Dung Ho My |last2=Panthum |first2=Thitipong |last3=Ponjarat |first3=Jatupong |last4=Laopichienpong |first4=Nararat |last5=Kraichak |first5=Ekaphan |last6=Singchat |first6=Worapong |last7=Ahmad |first7=Syed Farhan |last8=Muangmai |first8=Narongrit |last9=Peyachoknagul |first9=Surin |last10=Na-Nakorn |first10=Uthairat |last11=Srikulnath |first11=Kornsorn |date=2021-01-05 |title=An Investigation of ZZ/ZW and XX/XY Sex Determination Systems in North African Catfish (Clarias gariepinus, Burchell, 1822) |journal=Frontiers in Genetics |volume=11 |doi=10.3389/fgene.2020.562856 |doi-access=free |pmid=33584785 |issn=1664-8021|pmc=7874028 }}

Multiple sex chromosomes like those of platypus also occurs in bony fish.{{Cite journal |last1=Sember |first1=Alexandr |last2=Nguyen |first2=Petr |last3=Perez |first3=Manolo F. |last4=Altmanová |first4=Marie |last5=Ráb |first5=Petr |last6=Cioffi |first6=Marcelo de Bello |date=2021-09-13 |title=Multiple sex chromosomes in teleost fishes from a cytogenetic perspective: state of the art and future challenges |journal=Philosophical Transactions of the Royal Society B: Biological Sciences |volume=376 |issue=1833 |pages=20200098 |doi=10.1098/rstb.2020.0098 |issn=0962-8436 |pmc=8310710 |pmid=34304595}} Some moths and butterflies have $W\_1W\_2Z\; \backslash text\; \{♀\}/ZZ\; \backslash text\{♂\}$ or $WZ\_1Z\_2\; \backslash text\; \{♀\}/Z\_1Z\_1Z\_2Z\_2\; \backslash text\{♂\}$.{{Cite journal |last1=Traut |first1=W. |last2=Sahara |first2=K. |last3=Marec |first3=F. |date=2008-01-18 |title=Sex Chromosomes and Sex Determination in Lepidoptera |url=https://doi.org/10.1159/000111765 |journal=Sexual Development |volume=1 |issue=6 |pages=332–346 |doi=10.1159/000111765 |pmid=18391545 |issn=1661-5425}}

The Southern platyfish has a complex sex determination system involving 3 sex chromosomes and 4 autosomal alleles.{{Citation |last=Kallman |first=Klaus D. |title=A New Look at Sex Determination in Poeciliid Fishes |date=1984 |work=Evolutionary Genetics of Fishes |pages=95–171 |editor-last=Turner |editor-first=Bruce J. |url=https://doi.org/10.1007/978-1-4684-4652-4_3 |access-date=2024-08-04 |place=Boston, MA |publisher=Springer US |doi=10.1007/978-1-4684-4652-4_3 |isbn=978-1-4684-4652-4}}{{cite journal |vauthors=Schartl M |date=July 2004 |title=A comparative view on sex determination in medaka |journal=Mechanisms of Development |volume=121 |issue=7–8 |pages=639–645 |doi=10.1016/j.mod.2004.03.001 |pmid=15210173 |s2cid=17401686 |doi-access=free}}

Gastrotheca pseustes has $XY\_b\; \backslash text\; \{♀\}\; /\; XY\_a\; \backslash text\{♂\}$ C-banding heteromorphism, meaning that both males and females have XY chromosomes, but their Y chromosomes are different on one or more C-bands. Eleutherodactylus maussi has a $\backslash mathrm\{X\}\_1\; \backslash mathrm\{X\}\_1\; \backslash mathrm\{X\}\_2\; \backslash mathrm\{X\}\_2\; \backslash text\; \{♀\}\; /\; \backslash mathrm\{X\}\_1\; \backslash mathrm\{X\}\_2\; \backslash mathrm\{Y\}\; \backslash text\{♂\}$ system.{{Cite journal |last1=Schmid |first1=M. |last2=Steinlein |first2=C. |last3=Feichtinger |first3=W. |date=March 1992 |title=Chromosome banding in amphibia: XVII. First demonstration of multiple sex chromosomes in amphibians: Eleutherodactylus maussi (Anura, Leptodactylidae) |url=http://link.springer.com/10.1007/BF00346007 |journal=Chromosoma |volume=101 |issue=5–6 |pages=284–292 |doi=10.1007/BF00346007 |pmid=1576881 |issn=0009-5915}}{{Cite journal |last1=Schmid |first1=M. |last2=Feichtinger |first2=W. |last3=Steinlein |first3=C. |last4=Haaf |first4=T. |last5=Schartl |first5=M. |last6=Visbal García |first6=R. |last7=Manzanilla Pupo |first7=J. |last8=Fernández Badillo |first8=A. |date=2003-08-14 |title=Chromosome banding in Amphibia: XXVI. Coexistence of homomorphic XY sex chromosomes and a derived Y-autosome translocation in Eleutherodactylus maussi (Anura, Leptodactylidae) |url=https://doi.org/10.1159/000071612 |journal=Cytogenetics and Cell Genetics |volume=99 |issue=1–4 |pages=330–343 |doi=10.1159/000071612 |pmid=12900583 |issn=0301-0171}}

See {{Cite journal |last1=Zhu |first1=Zexian |last2=Younas |first2=Lubna |last3=Zhou |first3=Qi |date=2024-07-18 |title=Evolution and regulation of animal sex chromosomes |url=https://www.nature.com/articles/s41576-024-00757-3 |journal=Nature Reviews Genetics |pages=1–16 |doi=10.1038/s41576-024-00757-3 |pmid=39026082 |issn=1471-0064}} for a review.

Sexual chromosome pairs can arise from an autosomal pair that, for various reasons, stopped recombination, allowing for their divergence. The rate at which recombination is suppressed, and therefore the rate of sex chromosome divergence, is very different across clades.

In analogy with geological strata, historical events in the evolution of sex chromosomes are called evolutionary strata. The human Y-chromosome has had about 5 strata since the origin of the X and Y chromosomes about 300 Mya from a pair of autosomes. Each stratum was formed when a pseudoautosomal region (PAR) of the Y chromosome is inverted, stopping it from recombination with the X chromosome. Over time, each inverted region decays, possibly due to Muller's ratchet.{{Cite journal |last1=Lahn |first1=Bruce T. |last2=Page |first2=David C. |date=1999-10-29 |title=Four Evolutionary Strata on the Human X Chromosome |url=https://www.science.org/doi/10.1126/science.286.5441.964 |journal=Science |volume=286 |issue=5441 |pages=964–967 |doi=10.1126/science.286.5441.964 |pmid=10542153 |issn=0036-8075}}{{Cite journal |last1=Lemaitre |first1=Claire |last2=Braga |first2=Marilia D. V. |last3=Gautier |first3=Christian |last4=Sagot |first4=Marie-France |last5=Tannier |first5=Eric |last6=Marais |first6=Gabriel A. B. |date=2009-01-01 |title=Footprints of Inversions at Present and Past Pseudoautosomal Boundaries in Human Sex Chromosomes |journal=Genome Biology and Evolution |volume=1 |pages=56–66 |doi=10.1093/gbe/evp006 |issn=1759-6653 |pmc=2817401 |pmid=20333177}} Primate Y-chromosome evolution was rapid, with multiple inversions and shifts of the boundary of PAR.{{Cite journal |last1=Zhou |first1=Yang |last2=Zhan |first2=Xiaoyu |last3=Jin |first3=Jiazheng |last4=Zhou |first4=Long |last5=Bergman |first5=Juraj |last6=Li |first6=Xuemei |last7=Rousselle |first7=Marjolaine Marie C. |last8=Belles |first8=Meritxell Riera |last9=Zhao |first9=Lan |last10=Fang |first10=Miaoquan |last11=Chen |first11=Jiawei |last12=Fang |first12=Qi |last13=Kuderna |first13=Lukas |last14=Marques-Bonet |first14=Tomas |last15=Kitayama |first15=Haruka |date=July 2023 |title=Eighty million years of rapid evolution of the primate Y chromosome |url=https://www.nature.com/articles/s41559-022-01974-x |journal=Nature Ecology & Evolution |volume=7 |issue=7 |pages=1114–1130 |doi=10.1038/s41559-022-01974-x |pmid=37268856 |bibcode=2023NatEE...7.1114Z |issn=2397-334X}}

Among many species of the salamanders, the two chromosomes are only distinguished by a pericentric inversion, so that the banding pattern of the X chromosome is the same as that of Y, but with a region near the centromere reversed. (fig 7 {{Cite book |last=Solari |first=Alberto J. |title=Sex chromosomes and sex determination in vertebrates |date=1994 |publisher=CRC Press |isbn=978-0-8493-4571-5 |location=Boca Raton}}) In some species, the X is pericentrically inverted and the Y is ancestral. In other species it is the opposite. (p. 15 )

The gene content of the X chromosome is almost identical among placental mammals. This is hypothesized to be because the X inactivation means any change would cause serious disruption, thus subjecting it to strong purifying selection. Similarly, birds have highly conserved Z chromosomes.

Neo-sex chromosomes are currently existing sex chromosomes that formed when an autosome pair fused to the previously existing sex chromosome pair. Following this fusion, the autosomal portion undergoes recombination suppression, allowing them to differentiate. Such systems have been observed in insects, reptiles, birds, and mammals. They are useful to the study of the evolution of Y chromosome degeneration and dosage compensation.{{Cite journal |last1=Zhou |first1=Qi |last2=Wang |first2=Jun |last3=Huang |first3=Ling |last4=Nie |first4=Wenhui |last5=Wang |first5=Jinhuan |last6=Liu |first6=Yan |last7=Zhao |first7=Xiangyi |last8=Yang |first8=Fengtang |last9=Wang |first9=Wen |date=2008-06-14 |title=Neo-sex chromosomes in the black muntjac recapitulate incipient evolution of mammalian sex chromosomes |journal=Genome Biology |volume=9 |issue=6 |pages=R98 |doi=10.1186/gb-2008-9-6-r98 |doi-access=free |issn=1474-760X |pmc=2481430 |pmid=18554412}}{{Cite journal |last1=Pala |first1=I. |last2=Naurin |first2=S. |last3=Stervander |first3=M. |last4=Hasselquist |first4=D. |last5=Bensch |first5=S. |last6=Hansson |first6=B. |date=March 2012 |title=Evidence of a neo-sex chromosome in birds |journal=Heredity |volume=108 |issue=3 |pages=264–272 |doi=10.1038/hdy.2011.70 |pmid=21897438 |pmc=3282394 |issn=1365-2540}}

The sex-chromosome turnover is an evolutionary phenomenon where sex chromosomes disappear, or becomes autosomal, and autosomal chromosomes become sexual, repeatedly over evolutionary time. Some lineages have extensive turnover, but others don't. Generally, in an XY system, if the Y chromosome is degenerate, mostly different from the X chromosome, and has X dosage compensation, then turnover is unlikely. In particular, this applies to humans.{{Cite journal |last=Vicoso |first=Beatriz |date=December 2019 |title=Molecular and evolutionary dynamics of animal sex-chromosome turnover |url=https://www.nature.com/articles/s41559-019-1050-8 |journal=Nature Ecology & Evolution |volume=3 |issue=12 |pages=1632–1641 |doi=10.1038/s41559-019-1050-8 |bibcode=2019NatEE...3.1632V |issn=2397-334X}}{{Cite journal |last1=Palmer |first1=Daniela H. |last2=Rogers |first2=Thea F. |last3=Dean |first3=Rebecca |last4=Wright |first4=Alison E. |date=November 2019 |title=How to identify sex chromosomes and their turnover |journal=Molecular Ecology |volume=28 |issue=21 |pages=4709–4724 |doi=10.1111/mec.15245 |issn=0962-1083 |pmc=6900093 |pmid=31538682|bibcode=2019MolEc..28.4709P }}

The ZW and XY systems can evolve into to each other due to sexual conflict.{{Cite journal |last1=van Doorn |first1=G Sander |last2=Kirkpatrick |first2=Mark |date=2010-10-01 |title=Transitions Between Male and Female Heterogamety Caused by Sex-Antagonistic Selection |journal=Genetics |volume=186 |issue=2 |pages=629–645 |doi=10.1534/genetics.110.118596 |issn=1943-2631 |pmc=2954476 |pmid=20628036}}

It is an evolutionary puzzle why certain sex chromosomes remain homomorphic over millions of years, especially among lineages of fishes, amphibians, and nonavian reptiles. The fountain-of-youth model states that heteromorphy results from recombination suppression, and recombination suppression results from the male phenotype, not the sex chromosomes themselves. Therefore, if some XY sex-reversed females are fertile and adaptive under some circumstances, then the X and Y chromosomes would recombine in these individuals, preventing Y chromosome decay and maintaining long-term homomorphism.{{Cite journal |last=Perrin |first=Nicolas |date=December 2009 |title=Sex Reversal: A Fountain of Youth for Sex Chromosomes? |url=https://academic.oup.com/evolut/article/63/12/3043/6881025 |journal=Evolution |volume=63 |issue=12 |pages=3043–3049 |doi=10.1111/j.1558-5646.2009.00837.x|pmid=19744117 }}

Sex reversal denotes a situation where the phenotypic sex is different from the genotypic sex. While in humans, sex reversal (such as the XX male syndrome) are often infertile, sex-reversed individuals of some species are fertile under some conditions. For example, some XY-individuals in population of Chinook salmon in the Columbia River became fertile females, producing YY sons. Since Chinook salmons have homomorphic sex chromosomes, such YY sons are healthy. When YY males mate with XX females, all their progeny would be XY male if grown under normal conditions.{{Cite journal |last1=Nagler |first1=J J |last2=Bouma |first2=J |last3=Thorgaard |first3=G H |last4=Dauble |first4=D D |date=January 2001 |title=High incidence of a male-specific genetic marker in phenotypic female chinook salmon from the Columbia River. |journal=Environmental Health Perspectives |volume=109 |issue=1 |pages=67–69 |doi=10.1289/ehp.0110967 |issn=0091-6765 |pmc=1242053 |pmid=11171527}}

Support for the hypothesis is found in the common frog, for which XX males and XY males both suppresses sex chromosome recombination, but XX and XY females both recombine at the same rate.{{Cite journal |last1=Rodrigues |first1=Nicolas |last2=Studer |first2=Tania |last3=Dufresnes |first3=Christophe |last4=Perrin |first4=Nicolas |date=2018-04-01 |title=Sex-Chromosome Recombination in Common Frogs Brings Water to the Fountain-of-Youth |url=https://academic.oup.com/mbe/article/35/4/942/4823133 |journal=Molecular Biology and Evolution |volume=35 |issue=4 |pages=942–948 |doi=10.1093/molbev/msy008 |issn=0737-4038}}

{{main |Environmental sex determination}}

{{Main |Temperature-dependent sex determination}}

File:Alligator.jpg

Many other sex-determination systems exist. In some species of reptiles, including alligators, some turtles, and the tuatara, sex is determined by the temperature at which the egg is incubated during a temperature-sensitive period. There are no examples of temperature-dependent sex determination (TSD) in birds. Megapodes had formerly been thought to exhibit this phenomenon, but were found to actually have different temperature-dependent embryo mortality rates for each sex.{{cite journal | vauthors = Göth A, Booth DT | title = Temperature-dependent sex ratio in a bird | journal = Biology Letters | volume = 1 | issue = 1 | pages = 31–33 | date = March 2005 | pmid = 17148121 | pmc = 1629050 | doi = 10.1098/rsbl.2004.0247 }} For some species with TSD, sex determination is achieved by exposure to hotter temperatures resulting in the offspring being one sex and cooler temperatures resulting in the other. This type of TSD is called Pattern I. For others species using TSD, it is exposure to temperatures on both extremes that results in offspring of one sex, and exposure to moderate temperatures that results in offspring of the opposite sex, called Pattern II TSD. The specific temperatures required to produce each sex are known as the female-promoting temperature and the male-promoting temperature.{{cite journal | vauthors = Torres Maldonado LC, Landa Piedra A, Moreno Mendoza N, Marmolejo Valencia A, Meza Martínez A, Merchant Larios H | title = Expression profiles of Dax1, Dmrt1, and Sox9 during temperature sex determination in gonads of the sea turtle Lepidochelys olivacea | journal = General and Comparative Endocrinology | volume = 129 | issue = 1 | pages = 20–26 | date = October 2002 | pmid = 12409092 | doi = 10.1016/s0016-6480(02)00511-7 }} When the temperature stays near the threshold during the temperature sensitive period, the sex ratio is varied between the two sexes.{{cite journal | vauthors = Bull JJ |title=Sex Determination in Reptiles |journal=The Quarterly Review of Biology |date=March 1980 |volume=55 |issue=1 |pages=3–21 |jstor=2826077 |doi=10.1086/411613|s2cid=85177125 }} Some species' temperature standards are based on when a particular enzyme is created. These species that rely upon temperature for their sex determination do not have the SRY gene, but have other genes such as DAX1, DMRT1, and SOX9 that are expressed or not expressed depending on the temperature. The sex of some species, such as the Nile tilapia, Australian skink lizard, and Australian dragon lizard, has an initial bias, set by chromosomes, but can later be changed by the temperature of incubation.

It is unknown how exactly temperature-dependent sex determination evolved.{{cite journal |vauthors=Valenzuela N, Janzen FJ |title=Nest-site philopatry and the evolution of temperature-dependent sex determination |journal=Evolutionary Ecology Research |year=2001 |volume=3 |pages=779–794 |url=http://www.public.iastate.edu/~fjanzen/pdf/01EvolEcolRes.pdf |access-date=7 December 2011 |archive-date=4 May 2013 |archive-url=https://web.archive.org/web/20130504051408/http://www.public.iastate.edu/~fjanzen/pdf/01EvolEcolRes.pdf |url-status=live }} It could have evolved through certain sexes being more suited to certain areas that fit the temperature requirements. For example, a warmer area could be more suitable for nesting, so more females are produced to increase the amount that nest next season.

In amniotes, environmental sex determination preceded the genetically determined systems of birds and mammals; it is thought that a temperature-dependent amniote was the common ancestor of amniotes with sex chromosomes.{{cite journal | vauthors = Janzen FJ, Phillips PC | title = Exploring the evolution of environmental sex determination, especially in reptiles | journal = Journal of Evolutionary Biology | volume = 19 | issue = 6 | pages = 1775–1784 | date = November 2006 | pmid = 17040374 | doi = 10.1111/j.1420-9101.2006.01138.x | s2cid = 15485510 }}

There are other environmental sex determination systems including location-dependent determination systems as seen in the marine worm Bonellia viridis – larvae become males if they make physical contact with a female, and females if they end up on the bare sea floor. This is triggered by the presence of a chemical produced by the females, bonellin.{{cite book | vauthors = Gilbert S |title=Developmental biology |url=https://archive.org/details/developmentalbio00gilb_292 |url-access=limited |date=2006 |publisher=Sinauer Associates, Inc. Publishers |location=Sunderland, Mass. |isbn=9780878932504 |pages=[https://archive.org/details/developmentalbio00gilb_292/page/n569 550]–553 |edition= 8th.}} Some species, such as some snails, practice sex change: adults start out male, then become female. In tropical clownfish, the dominant individual in a group becomes female while the other ones are male, and bluehead wrasses (Thalassoma bifasciatum) are the reverse.

File:ClownFishCycle.jpg

Clownfish live in colonies of several small undifferentiated fish and two large fish (male and female). The male and female are the only sexually mature fish to reproduce. Clownfish are protandrous hermaphrodites, which means after they mature into males, they eventually can transform into females. They develop undifferentiated until they are needed to fill a certain role in their environment, i.e., if they receive the social and environmental cues to do so. Casas, L., Saborido-Rey, F., Ryu, T., Michell, C., Ravasi, T., & Irigoien, X. (2016). Sex Change in Clownfish: Molecular Insights from Transcriptome Analysis. Scientific Reports, 6(1). https://doi.org/10.1038/srep35461

Some species, however, have no sex-determination system. Hermaphrodite species include the common earthworm and certain species of snails. A few species of fish, reptiles, and insects reproduce by parthenogenesis and are female altogether. There are some reptiles, such as the boa constrictor and Komodo dragon that can reproduce both sexually and asexually, depending on whether a mate is available.{{cite journal | vauthors = Watts PC, Buley KR, Sanderson S, Boardman W, Ciofi C, Gibson R | title = Parthenogenesis in Komodo dragons | journal = Nature | volume = 444 | issue = 7122 | pages = 1021–1022 | date = December 2006 | pmid = 17183308 | doi = 10.1038/4441021a | name-list-style = amp | s2cid = 4311088 | bibcode = 2006Natur.444.1021W }}

There are exceptional sex-determination systems, neither genetic nor environmental.

The Wolbachia genus of parasitic bacteria lives inside the cytoplasm of its host, and is vertically transmitted from parents to children. They primarily infect arthropods and nematodes. Different Wolbachia can alter the reproductive abilities of its host by a variety of means, including cytoplasmic incompatibility, parthenogenesis, feminization and embryonic male killing.{{Cite journal |last1=Stevens |first1=Lori |last2=Giordano |first2=Rosanna |last3=Fialho |first3=Roberto F. |date=November 2001 |title=Male-Killing, Nematode Infections, Bacteriophage Infection, and Virulence of Cytoplasmic Bacteria in the Genus Wolbachia |url=https://www.annualreviews.org/doi/10.1146/annurev.ecolsys.32.081501.114132 |journal=Annual Review of Ecology and Systematics |language=en |volume=32 |issue=1 |pages=519–545 |doi=10.1146/annurev.ecolsys.32.081501.114132 |issn=0066-4162}}

Mitochondrial male sterility: In many flowering plants, the mitochondria can cause hermaphrodite individuals to be unable to father offspring, effectively turning them into exclusive females. This is a form of mother’s curse. It is an evolutionarily adaptive strategy for mitochondria as mitochondria are inherited exclusively from mother to offspring. The first published case of mitochondrial male sterility among metazoans was reported in 2022 in the hermaphroditic snail Physa acuta.{{Cite journal |last1=David |first1=Patrice |last2=Degletagne |first2=Cyril |last3=Saclier |first3=Nathanaëlle |last4=Jennan |first4=Aurel |last5=Jarne |first5=Philippe |last6=Plénet |first6=Sandrine |last7=Konecny |first7=Lara |last8=François |first8=Clémentine |last9=Guéguen |first9=Laurent |last10=Garcia |first10=Noéline |last11=Lefébure |first11=Tristan |last12=Luquet |first12=Emilien |date=May 2022 |title=Extreme mitochondrial DNA divergence underlies genetic conflict over sex determination |url=https://linkinghub.elsevier.com/retrieve/pii/S0960982222005838 |journal=Current Biology |language=en |volume=32 |issue=10 |pages=2325–2333.e6 |doi=10.1016/j.cub.2022.04.014}}

In some species of insects, springtails and mites, male offspring lose their paternal genome (in whole or in part) during development or in the germline. Males can either be diploid, diploid with missing sex chromosome, functionally haploid or truly haploid, depending on the mechanism of elimination.{{Cite journal |last=Herbette |first=Marion |last2=Ross |first2=Laura |date=2023 |title=Paternal genome elimination: patterns and mechanisms of drive and silencing |url=https://linkinghub.elsevier.com/retrieve/pii/S0959437X2300045X |journal=Current Opinion in Genetics & Development |language=en |volume=81 |pages=102065 |doi=10.1016/j.gde.2023.102065|doi-access=free }}

In some species of Hymenoptera (ants, bees and wasps), flies and crustaceans, all offspring of a particular individual female are either exclusively male or exclusively female. The underlying mechanisms are diverse and include maternally controlled paternal genome elimination and Mendelian inherited maternal sex-determining factors.{{Cite journal |last=Baird |first=Robert B. |last2=Mongue |first2=Andrew J. |last3=Ross |first3=Laura |date=2023 |title=Why put all your eggs in one basket? Evolutionary perspectives on the origins of monogenic reproduction |url=https://www.nature.com/articles/s41437-023-00632-7 |journal=Heredity |language=en |volume=131 |issue=2 |pages=87–95 |doi=10.1038/s41437-023-00632-7 |issn=0018-067X |pmc=10382564 |pmid=37328587}}

File:A region in the pseudoautosomal region of the short arms of the X- and Y-chromosome.jpg , highlighted here in green, are all that is left of the original autosomes that can still cross over with each other.]]

Sex determination systems may have evolved from mating type, which is a feature of microorganisms.

Chromosomal sex determination may have evolved early in the history of eukaryotes.{{cite journal | vauthors = Lehtonen J, Parker GA | title = Gamete competition, gamete limitation, and the evolution of the two sexes | journal = Molecular Human Reproduction | volume = 20 | issue = 12 | pages = 1161–1168 | date = December 2014 | pmid = 25323972 | doi = 10.1093/molehr/gau068 | doi-access = free }} But in plants it has been suggested to have evolved recently.{{Cite book | vauthors = Clarke R, Merlin M |url=https://books.google.com/books?id=bs4hEAAAQBAJ&q=XY+sex+determination+in+plants&pg=PA359 |title=Cannabis: Evolution and Ethnobotany |date=2016-06-28 |publisher=Univ of California Press |isbn=978-0-520-29248-2 |pages=359 }}

The accepted hypothesis of XY and ZW sex chromosome evolution in amniotes is that they evolved at the same time, in two different branches.{{cite journal | vauthors = Namekawa SH, Lee JT | title = XY and ZW: is meiotic sex chromosome inactivation the rule in evolution? | journal = PLOS Genetics | volume = 5 | issue = 5 | pages = e1000493 | date = May 2009 | pmid = 19461890 | pmc = 2679206 | doi = 10.1371/journal.pgen.1000493 | doi-access = free }}{{cite journal | vauthors = Vallender EJ, Lahn BT | title = Multiple independent origins of sex chromosomes in amniotes | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 103 | issue = 48 | pages = 18031–18032 | date = November 2006 | pmid = 17116892 | pmc = 1838700 | doi = 10.1073/pnas.0608879103 | doi-access = free | bibcode = 2006PNAS..10318031V }}

No genes are shared between the avian ZW and mammal XY chromosomes and the chicken Z chromosome is similar to the human autosomal chromosome 9, rather than X or Y. This suggests not that the ZW and XY sex-determination systems share an origin but that the sex chromosomes are derived from autosomal chromosomes of the common ancestor of birds and mammals. In the platypus, a monotreme, the X₁ chromosome shares homology with therian mammals, while the X₅ chromosome contains an avian sex-determination gene, further suggesting an evolutionary link.{{cite journal | vauthors = Veyrunes F, Waters PD, Miethke P, Rens W, McMillan D, Alsop AE, Grützner F, Deakin JE, Whittington CM, Schatzkamer K, Kremitzki CL, Graves T, Ferguson-Smith MA, Warren W, Marshall Graves JA | display-authors = 6 | title = Bird-like sex chromosomes of platypus imply recent origin of mammal sex chromosomes | journal = Genome Research | volume = 18 | issue = 6 | pages = 965–973 | date = June 2008 | pmid = 18463302 | pmc = 2413164 | doi = 10.1101/gr.7101908 }}

However, there is some evidence to suggest that there could have been transitions between ZW and XY, such as in Xiphophorus maculatus, which have both ZW and XY systems in the same population, despite the fact that ZW and XY have different gene locations.{{cite journal | vauthors = Marshall Graves JA | title = Human Y chromosome, sex determination, and spermatogenesis- a feminist view | journal = Biology of Reproduction | volume = 63 | issue = 3 | pages = 667–676 | date = September 2000 | pmid = 10952906 | doi = 10.1095/biolreprod63.3.667b | doi-access = free }}{{cite journal | vauthors = Ezaz T, Stiglec R, Veyrunes F, Marshall Graves JA | title = Relationships between vertebrate ZW and XY sex chromosome systems | journal = Current Biology | volume = 16 | issue = 17 | pages = R736–R743 | date = September 2006 | pmid = 16950100 | doi = 10.1016/j.cub.2006.08.021 | bibcode = 2006CBio...16.R736E | hdl-access = free | s2cid = 18864471 | hdl = 1885/37887 }} A recent theoretical model raises the possibility of both transitions between the XY/XX and ZZ/ZW system and environmental sex determination{{cite journal | vauthors = Quinn AE, Sarre SD, Ezaz T, Marshall Graves JA, Georges A | title = Evolutionary transitions between mechanisms of sex determination in vertebrates | journal = Biology Letters | volume = 7 | issue = 3 | pages = 443–448 | date = June 2011 | pmid = 21212104 | pmc = 3097877 | doi = 10.1098/rsbl.2010.1126 }} The platypus' genes also back up the possible evolutionary link between XY and ZW, because they have the DMRT1 gene possessed by birds on their X chromosomes. Regardless, XY and ZW follow a similar route. All sex chromosomes started out as an original autosome of an original amniote that relied upon temperature to determine the sex of offspring. After the mammals separated, the reptile branch further split into Lepidosauria and Archosauromorpha. These two groups both evolved the ZW system separately, as evidenced by the existence of different sex chromosomal locations. In mammals, one of the autosome pair, now Y, mutated its SOX3 gene into the SRY gene, causing that chromosome to designate sex.{{cite journal | vauthors = Graves JA | title = Sex chromosome specialization and degeneration in mammals | journal = Cell | volume = 124 | issue = 5 | pages = 901–914 | date = March 2006 | pmid = 16530039 | doi = 10.1016/j.cell.2006.02.024 | s2cid = 8379688 | doi-access = free }}{{cite press release | title=The evolution of the sex chromosomes: Step by step | publisher=University of Chicago Medical Center |date=28 October 1999 |url =http://www.uchospitals.edu/news/1999/19991028-x-vs-y.html |access-date=23 October 2011}} After this mutation, the SRY-containing chromosome inverted and was no longer completely homologous with its partner. The regions of the X and Y chromosomes that are still homologous to one another are known as the pseudoautosomal region.{{cite journal | vauthors = Charlesworth B | title = The organization and evolution of the human Y chromosome | journal = Genome Biology | volume = 4 | issue = 9 | pages = 226 | date = 14 August 2003 | pmid = 12952526 | pmc = 193647 | doi = 10.1186/gb-2003-4-9-226 | doi-access = free }} Once it inverted, the Y chromosome became unable to remedy deleterious mutations, and thus degenerated. There is some concern that the Y chromosome will shrink further and stop functioning in ten million years: but the Y chromosome has been strictly conserved after its initial rapid gene loss.{{cite journal | vauthors = Graves JA | title = The degenerate Y chromosome--can conversion save it? | journal = Reproduction, Fertility, and Development | volume = 16 | issue = 5 | pages = 527–534 | date = 22 July 2004 | pmid = 15367368 | doi = 10.1071/RD03096 | s2cid = 23740483 }}{{cite journal | vauthors = Hughes JF, Skaletsky H, Brown LG, Pyntikova T, Graves T, Fulton RS, Dugan S, Ding Y, Buhay CJ, Kremitzki C, Wang Q, Shen H, Holder M, Villasana D, Nazareth LV, Cree A, Courtney L, Veizer J, Kotkiewicz H, Cho TJ, Koutseva N, Rozen S, Muzny DM, Warren WC, Gibbs RA, Wilson RK, Page DC | display-authors = 6 | title = Strict evolutionary conservation followed rapid gene loss on human and rhesus Y chromosomes | journal = Nature | volume = 483 | issue = 7387 | pages = 82–86 | date = February 2012 | pmid = 22367542 | pmc = 3292678 | doi = 10.1038/nature10843 | bibcode = 2012Natur.483...82H }}

There are some vertebrate species, such as the medaka fish, that evolved sex chromosomes separately; their Y chromosome never inverted and can still swap genes with the X. These species' sex chromosomes are relatively primitive and unspecialized. Because the Y does not have male-specific genes and can interact with the X, XY and YY females can be formed as well as XX males. Non-inverted Y chromosomes with long histories are found in pythons and emus, each system being more than 120 million years old, suggesting that inversions are not necessarily an eventuality.{{cite journal | vauthors = Bachtrog D, Mank JE, Peichel CL, Kirkpatrick M, Otto SP, Ashman TL, Hahn MW, Kitano J, Mayrose I, Ming R, Perrin N, Ross L, Valenzuela N, Vamosi JC | display-authors = 6 | title = Sex determination: why so many ways of doing it? | journal = PLOS Biology | volume = 12 | issue = 7 | pages = e1001899 | date = July 2014 | pmid = 24983465 | pmc = 4077654 | doi = 10.1371/journal.pbio.1001899 | doi-access = free }} XO sex determination can evolve from XY sex determination with about 2 million years.{{clarification needed|reason=What does this sentence mean?|date=April 2022}}{{Cite book| vauthors = Nei M |url=https://books.google.com/books?id=sJtoAgAAQBAJ&q=sexual+systems+can+evolve&pg=PA168|title=Mutation-Driven Evolution|date=2013-05-02|publisher=OUP Oxford|isbn=978-0-19-163781-0|pages=168}}

• Clarence Erwin McClung, who discovered the role of chromosomes in sex determination
• Testis-determining factor
• Maternal influence on sex determination
• Sequential hermaphroditism
• Sex determination and differentiation (human)
• Cell autonomous sex identity

{{Reflist}}

{{refbegin}}

• {{cite book |vauthors=Beukeboom L, Perrin N |date=2014 |url=https://books.google.com/books?id=d4cLBAAAQBAJ |title=The Evolution of Sex Determination |publisher=Oxford University Press |isbn=978-0-19-163140-5}}
• {{Cite paper |last=Lahn |first=Bruce T. |date=1999-10-29 |title=Four Evolutionary Strata on the Human X Chromosome |url=https://www.science.org/doi/10.1126/science.286.5441.964 |journal=Science |volume=286 |issue=5441 |pages=964–967 |doi=10.1126/science.286.5441.964 |issn=0036-8075 |pmid=10542153}}

{{refend}}

{{Sex determination and differentiation}}

{{Sex (biology)}}

{{chromo}}

{{Authority control}}

{{DEFAULTSORT:Sex-Determination System}}

Category:Epigenetics