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Haldane's rule

April 03, 2024

Not to be confused with Haldane's principle.

Haldane's rule is an observation about the early stage of speciation, formulated in 1922 by the British evolutionary biologist J. B. S. Haldane, that states that if — in a species hybrid — only one sex is inviable or sterile, that sex is more likely to be the heterogametic sex. The heterogametic sex is the one with two different sex chromosomes; in therian mammals,[a] for example, this is the male.[2]

In humans, barring intersex conditions causing aneuploidy and other unusual states, it is the male that is heterogametic, with XY sex chromosomes.

Overview edit

Haldane himself described the rule as:

When in the F1 offspring of two different animal races one sex is absent, rare, or sterile, that sex is the heterozygous sex (heterogametic sex).[3]

Haldane's rule applies to the vast majority of heterogametic organisms. This includes the case where two species make secondary contact in an area of sympatry and form hybrids after allopatric speciation has occurred.

The rule includes both male heterogametic (XY or XO-type sex determination, such as found in mammals and Drosophila fruit flies) and female heterogametic (ZW or Z0-type sex determination, as found in birds and butterflies), and some dioecious plants such as campions.[4]

Hybrid dysfunction (sterility and inviability) is a major form of post-zygotic reproductive isolation, which occurs in early stages of speciation. Evolution can produce a similar pattern of isolation in a vast array of different organisms. However, the actual mechanisms leading to Haldane's rule in different taxa remain largely undefined.

Hypotheses edit

Many different hypotheses have been advanced to address the evolutionary mechanisms to produce Haldane's rule. Currently, the most popular explanation for Haldane's rule is the composite hypothesis, which divides Haldane's rule into multiple subdivisions, including sterility, inviability, male heterogamety, and female heterogamety. The composite hypothesis states that Haldane's rule in different subdivisions has different causes. Individual genetic mechanisms may not be mutually exclusive, and these mechanisms may act together to cause Haldane's rule in any given subdivision.[5][6] In contrast to these views that emphasize genetic mechanisms, another view hypothesizes that population dynamics during population divergence may cause Haldane's rule.[7]

The main genetic hypotheses are:

  • Dominance: Heterogametic hybrids are affected by all X-linked alleles (be they recessive or dominant) causing incompatibilities due to divergent alleles being brought together. However, homogametic hybrids are only affected by dominant deleterious X-linked alleles. Heterogametic hybrids, which carry only a single copy of a given X-linked gene, will be affected by mutations regardless of dominance. Thus, an X-linked incompatibility between diverging populations is more likely to be expressed in the heterogametic sex than in the homogametic sex.
  • The "faster male": Male genes are thought to evolve faster due to sexual selection.[6] As a result, male sterility becomes more evident in male heterogametic taxa (XY sex determination). This hypothesis conflicts with Haldane's rule in male homogametic taxa, in which females are more affected by hybrid inferiority. It therefore only applies to male sterility in taxa with XY sex determination, according to the composite theory.
  • Meiotic drive: In hybrid populations, selfish genetic elements inactivate sperm cells (i.e.: an X-linked drive factor inactivates a Y-bearing sperm and vice versa).
  • The "faster X": Genes on hemizygous chromosomes may evolve more quickly by enhancing selection on possible recessive alleles causing a larger effect in reproductive isolation.[8]
  • Differential selection: Hybrid incompatibilities affecting the heterogametic sex and homogametic sex are fundamentally different isolating mechanisms, which makes heterogametic inferiority (sterility/inviability) more visible or preserved in nature.[7]

Data from multiple phylogenetic groups support a combination of dominance and faster X-chromosome theories.[9] However, it has recently been argued that dominance theory can not explain Haldane's rule in marsupials since both sexes experience the same incompatibilities due to paternal X-inactivation in females.[10]

The dominance hypothesis is the core of the composite theory, and X-linked recessive/dominance effects have been demonstrated in many cases to cause hybrid incompatibilities. There is also supporting evidence for the faster male and meiotic drive hypotheses. For example, a significant reduction of male-driven gene flow is observed in Asian elephants, suggesting faster evolution of male traits.[11]

Although the rule was initially stated in context of diploid organisms with chromosomal sex determination, it has recently been argued that it can be extended to certain species lacking chromosomal sex determination, such as haplodiploids[12] and hermaphrodites.[9]

Exceptions edit

In some instances, the homogametic sex turns out to be inviable while the heterogametic sex is viable and fertile. This is seen in some Drosophila fruit flies.[13][14]

Notes edit

  1. ^ Unlike other mammals, monotremes have more than two different sex chromosomes: The platypus has five pairs. Short-beaked echidnas have four pairs plus one female-only chromosome.[1]

References edit

  1. ^ Deakin, J. E.; Graves, J. A. M.; Rens, W. (2012). "The Evolution of Marsupial and Monotreme Chromosomes". Cytogenetic and Genome Research. 137 (2–4): 113–129. doi:10.1159/000339433. hdl:1885/64794. PMID 22777195.
  2. ^ Turelli, M.; Orr, H. A. (May 1995). "The dominance theory of Haldane's rule". Genetics. 140 (1): 389–402. doi:10.1093/genetics/140.1.389. PMC 1206564. PMID 7635302.
  3. ^ Haldane, J. B. S. (1922). "Sex ratio and unisexual sterility in hybrid animals". J. Genet. 12 (2): 101–109. doi:10.1007/BF02983075. S2CID 32459333.
  4. ^ Brothers, Amanda N.; Delph, Lynda F. (2010). "Haldane's rule is extended to plants with sex chromosomes". Evolution. 64 (12): 3643–3648. doi:10.1111/j.1558-5646.2010.01095.x. PMID 20681984. S2CID 27145478.
  5. ^ Orr, H. A. (1993). "Haldane's rule has multiple genetic causes". Nature. 361 (6412): 532–533. Bibcode:1993Natur.361..532O. doi:10.1038/361532a0. PMID 8429905. S2CID 4304828.
  6. ^ a b Wu, C.-I.; Davis, A. W. (1993). "Evolution of postmating reproductive isolation: The composite nature of Haldane's rule and its genetic bases". The American Naturalist. 142 (22): 187–212. doi:10.1086/285534. JSTOR 2462812. PMID 19425975. S2CID 35214550.
  7. ^ a b Wang, R. (2003). "Differential strength of sex-biased hybrid inferiority in impeding gene flow may be a cause of Haldane's rule". Journal of Evolutionary Biology. 16 (2): 353–361. doi:10.1046/j.1420-9101.2003.00528.x. PMID 14635874. S2CID 7127922.
  8. ^ Charlesworth, B.; Coyne, J. A.; Barton, N. H. (1987). "The relative rates of evolution of sex chromosomes and autosomes". The American Naturalist. 130 (1): 113–146. doi:10.1086/284701. JSTOR 2461884. S2CID 84357596.
  9. ^ a b Schilthuizen, M.; Giesbers, M. C.; Beukeboom, L. W. (2011). "Haldane's rule in the 21st century". Heredity. 107 (2): 95–102. doi:10.1038/hdy.2010.170. PMC 3178397. PMID 21224879.
  10. ^ Watson, E.; Demuth, J. (2012). "Haldane's rule in marsupials: What happens when both sexes are functionally hemizygous?". Journal of Heredity. 103 (3): 453–458. doi:10.1093/jhered/esr154. PMC 3331990. PMID 22378959.
  11. ^ Fickel, J.; Lieckfeldt, D.; Ratanakorn, P.; Pitra, C. (2007). "Distribution of haplotypes and microsatellite alleles among Asian elephants (Elephas maximus) in Thailand". European Journal of Wildlife Research. 53 (4): 298–303. doi:10.1007/s10344-007-0099-x. S2CID 25507884.
  12. ^ Koevoets, T.; Beukeboom, L. W. (2009). "Genetics of postzygotic isolation and Haldane's rule in haplodiploids". Heredity. 102 (1): 16–23. doi:10.1038/hdy.2008.44. PMID 18523445.
  13. ^ Sawamura, K. (1996). "Maternal effect as a cause of exceptions for Haldane's rule". Genetics. 143 (1): 609–611. doi:10.1093/genetics/143.1.609. PMC 1207293. PMID 8722809.
  14. ^ Ferree, Patrick M.; Barbash, Daniel A. (2009). "Species-specific heterochromatin prevents mitotic chromosome Segregation to Cause hybrid lethality in Drosophila". PLOS Biology. 7 (10): e1000234. doi:10.1371/journal.pbio.1000234. PMC 2760206. PMID 19859525.

Further reading edit

  • Coyne, J. A. (1985). "The genetic basis of Haldane's rule". Nature. 314 (6013): 736–738. Bibcode:1985Natur.314..736C. doi:10.1038/314736a0. PMID 3921852. S2CID 4309254.
  • Forsdyke, Donald (2005). . Archived from the original on 9 June 2011. Retrieved 11 October 2006.
  • Naisbit, Russell E.; Jiggins, Chris D.; Linares, Mauricio; Salazar, Camilo; Mallet, James (2002). "Hybrid sterility, Haldane's rule, and speciation in Heliconius cydno and H. melpomene". Genetics. 161 (4): 1517–1526. doi:10.1093/genetics/161.4.1517. PMC 1462209. PMID 12196397.
  • Coyne, J. A.; Charlesworth, B.; Orr, H. A. (1991). "Haldane's Rule Revisited". Evolution. 45 (7). Society for the Study of Evolution: 1710–1714. doi:10.1111/j.1558-5646.1991.tb02677.x. JSTOR 2409792. PMID 28564141.

April 03, 2024
haldane, rule, confused, with, haldane, principle, observation, about, early, stage, speciation, formulated, 1922, british, evolutionary, biologist, haldane, that, states, that, species, hybrid, only, inviable, sterile, that, more, likely, heterogametic, heter. Not to be confused with Haldane s principle Haldane s rule is an observation about the early stage of speciation formulated in 1922 by the British evolutionary biologist J B S Haldane that states that if in a species hybrid only one sex is inviable or sterile that sex is more likely to be the heterogametic sex The heterogametic sex is the one with two different sex chromosomes in therian mammals a for example this is the male 2 In humans barring intersex conditions causing aneuploidy and other unusual states it is the male that is heterogametic with XY sex chromosomes Contents 1 Overview 2 Hypotheses 3 Exceptions 4 Notes 5 References 6 Further readingOverview editHaldane himself described the rule as When in the F1 offspring of two different animal races one sex is absent rare or sterile that sex is the heterozygous sex heterogametic sex 3 Haldane s rule applies to the vast majority of heterogametic organisms This includes the case where two species make secondary contact in an area of sympatry and form hybrids after allopatric speciation has occurred The rule includes both male heterogametic XY or XO type sex determination such as found in mammals and Drosophila fruit flies and female heterogametic ZW or Z0 type sex determination as found in birds and butterflies and some dioecious plants such as campions 4 Hybrid dysfunction sterility and inviability is a major form of post zygotic reproductive isolation which occurs in early stages of speciation Evolution can produce a similar pattern of isolation in a vast array of different organisms However the actual mechanisms leading to Haldane s rule in different taxa remain largely undefined Hypotheses editMany different hypotheses have been advanced to address the evolutionary mechanisms to produce Haldane s rule Currently the most popular explanation for Haldane s rule is the composite hypothesis which divides Haldane s rule into multiple subdivisions including sterility inviability male heterogamety and female heterogamety The composite hypothesis states that Haldane s rule in different subdivisions has different causes Individual genetic mechanisms may not be mutually exclusive and these mechanisms may act together to cause Haldane s rule in any given subdivision 5 6 In contrast to these views that emphasize genetic mechanisms another view hypothesizes that population dynamics during population divergence may cause Haldane s rule 7 The main genetic hypotheses are Dominance Heterogametic hybrids are affected by all X linked alleles be they recessive or dominant causing incompatibilities due to divergent alleles being brought together However homogametic hybrids are only affected by dominant deleterious X linked alleles Heterogametic hybrids which carry only a single copy of a given X linked gene will be affected by mutations regardless of dominance Thus an X linked incompatibility between diverging populations is more likely to be expressed in the heterogametic sex than in the homogametic sex The faster male Male genes are thought to evolve faster due to sexual selection 6 As a result male sterility becomes more evident in male heterogametic taxa XY sex determination This hypothesis conflicts with Haldane s rule in male homogametic taxa in which females are more affected by hybrid inferiority It therefore only applies to male sterility in taxa with XY sex determination according to the composite theory Meiotic drive In hybrid populations selfish genetic elements inactivate sperm cells i e an X linked drive factor inactivates a Y bearing sperm and vice versa The faster X Genes on hemizygous chromosomes may evolve more quickly by enhancing selection on possible recessive alleles causing a larger effect in reproductive isolation 8 Differential selection Hybrid incompatibilities affecting the heterogametic sex and homogametic sex are fundamentally different isolating mechanisms which makes heterogametic inferiority sterility inviability more visible or preserved in nature 7 Data from multiple phylogenetic groups support a combination of dominance and faster X chromosome theories 9 However it has recently been argued that dominance theory can not explain Haldane s rule in marsupials since both sexes experience the same incompatibilities due to paternal X inactivation in females 10 The dominance hypothesis is the core of the composite theory and X linked recessive dominance effects have been demonstrated in many cases to cause hybrid incompatibilities There is also supporting evidence for the faster male and meiotic drive hypotheses For example a significant reduction of male driven gene flow is observed in Asian elephants suggesting faster evolution of male traits 11 Although the rule was initially stated in context of diploid organisms with chromosomal sex determination it has recently been argued that it can be extended to certain species lacking chromosomal sex determination such as haplodiploids 12 and hermaphrodites 9 Exceptions editIn some instances the homogametic sex turns out to be inviable while the heterogametic sex is viable and fertile This is seen in some Drosophila fruit flies 13 14 Notes edit Unlike other mammals monotremes have more than two different sex chromosomes The platypus has five pairs Short beaked echidnas have four pairs plus one female only chromosome 1 References edit Deakin J E Graves J A M Rens W 2012 The Evolution of Marsupial and Monotreme Chromosomes Cytogenetic and Genome Research 137 2 4 113 129 doi 10 1159 000339433 hdl 1885 64794 PMID 22777195 Turelli M Orr H A May 1995 The dominance theory of Haldane s rule Genetics 140 1 389 402 doi 10 1093 genetics 140 1 389 PMC 1206564 PMID 7635302 Haldane J B S 1922 Sex ratio and unisexual sterility in hybrid animals J Genet 12 2 101 109 doi 10 1007 BF02983075 S2CID 32459333 Brothers Amanda N Delph Lynda F 2010 Haldane s rule is extended to plants with sex chromosomes Evolution 64 12 3643 3648 doi 10 1111 j 1558 5646 2010 01095 x PMID 20681984 S2CID 27145478 Orr H A 1993 Haldane s rule has multiple genetic causes Nature 361 6412 532 533 Bibcode 1993Natur 361 532O doi 10 1038 361532a0 PMID 8429905 S2CID 4304828 a b Wu C I Davis A W 1993 Evolution of postmating reproductive isolation The composite nature of Haldane s rule and its genetic bases The American Naturalist 142 22 187 212 doi 10 1086 285534 JSTOR 2462812 PMID 19425975 S2CID 35214550 a b Wang R 2003 Differential strength of sex biased hybrid inferiority in impeding gene flow may be a cause of Haldane s rule Journal of Evolutionary Biology 16 2 353 361 doi 10 1046 j 1420 9101 2003 00528 x PMID 14635874 S2CID 7127922 Charlesworth B Coyne J A Barton N H 1987 The relative rates of evolution of sex chromosomes and autosomes The American Naturalist 130 1 113 146 doi 10 1086 284701 JSTOR 2461884 S2CID 84357596 a b Schilthuizen M Giesbers M C Beukeboom L W 2011 Haldane s rule in the 21st century Heredity 107 2 95 102 doi 10 1038 hdy 2010 170 PMC 3178397 PMID 21224879 Watson E Demuth J 2012 Haldane s rule in marsupials What happens when both sexes are functionally hemizygous Journal of Heredity 103 3 453 458 doi 10 1093 jhered esr154 PMC 3331990 PMID 22378959 Fickel J Lieckfeldt D Ratanakorn P Pitra C 2007 Distribution of haplotypes and microsatellite alleles among Asian elephants Elephas maximus in Thailand European Journal of Wildlife Research 53 4 298 303 doi 10 1007 s10344 007 0099 x S2CID 25507884 Koevoets T Beukeboom L W 2009 Genetics of postzygotic isolation and Haldane s rule in haplodiploids Heredity 102 1 16 23 doi 10 1038 hdy 2008 44 PMID 18523445 Sawamura K 1996 Maternal effect as a cause of exceptions for Haldane s rule Genetics 143 1 609 611 doi 10 1093 genetics 143 1 609 PMC 1207293 PMID 8722809 Ferree Patrick M Barbash Daniel A 2009 Species specific heterochromatin prevents mitotic chromosome Segregation to Cause hybrid lethality in Drosophila PLOS Biology 7 10 e1000234 doi 10 1371 journal pbio 1000234 PMC 2760206 PMID 19859525 Further reading editCoyne J A 1985 The genetic basis of Haldane s rule Nature 314 6013 736 738 Bibcode 1985Natur 314 736C doi 10 1038 314736a0 PMID 3921852 S2CID 4309254 Forsdyke Donald 2005 Haldane s rule Archived from the original on 9 June 2011 Retrieved 11 October 2006 Naisbit Russell E Jiggins Chris D Linares Mauricio Salazar Camilo Mallet James 2002 Hybrid sterility Haldane s rule and speciation in Heliconius cydno and H melpomene Genetics 161 4 1517 1526 doi 10 1093 genetics 161 4 1517 PMC 1462209 PMID 12196397 Coyne J A Charlesworth B Orr H A 1991 Haldane s Rule Revisited Evolution 45 7 Society for the Study of Evolution 1710 1714 doi 10 1111 j 1558 5646 1991 tb02677 x JSTOR 2409792 PMID 28564141 Retrieved from https en wikipedia org w index php title Haldane 27s rule amp oldid 1195873505, wikipedia, wiki, book, books, library,

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