「關係相容性的科學背後」

Jokiniemi A, Turunen T, Kohonen M et al. (2025) Female-mediated selective sperm activation may remodel major histocompatibility complex-based mate choice decisions in humans Heredity 134, 321–330.

*** This study reveals a paradox: women prefer the body odours of MHC-similar men, but sperm from MHC-dissimilar men exhibit higher motility when exposed to female follicular fluid. This suggests that the most attractive males may not necessarily be the most optimal partners in terms of fertilisation success, indicating that post-copulatory selection can remodel pre-mating mate choice decisions.

Havlíček J, Winternitz J, Roberts S (2020) Major histocompatibility complex-associated odour preferences and human mate choice: near and far horizons Philosophical Transactions of the Royal Society B, 375:20190260.

*** A comprehensive meta-analysis examining MHC-based mate choice in humans. The analysis found no significant overall effect of MHC-similarity on human mate selection when combining genomic studies, relationship satisfaction measures, and odour preference experiments. The authors recommend larger, more diverse samples and greater focus on mechanisms of MHC-associated pregnancy loss.

Dandine-Roulland C, Laurent R, Dall'Ara I, Toupance B 和 Chaix R (2019) 人类 MHC 不相配交配的基因组证据

This study on major histocompatibility complex (MHC) and mate choice in humans suggests that while couples in Northern Europe tend to exhibit MHC-dissimilarity, potentially indicative of a biological influence on mate choice, social and/or cultural factors in other regions, like Israel, may override or obscure such biologically-driven mating preferences.

Birnbaum GE, Zholtack K, Mizrahi M & Ein-Dor T (2019) 苦药：停用口服避孕药增强了人们的吸引力另类伴侣 Evolutionary Psychological Science volume 5, pages 276–285

*** 激素避孕藥改變了女性對伴侶的偏好，使其傾向於更有照顧能力的男性，但停止使用後，這些偏好會恢復，在高生育期時尤其會增強對更具遺傳相容性的男性的吸引力，可能會促使尋求替代伴侶。

Wedekind C (2019) A predicted interaction between odour pleasantness and intensity provides evidence for major histocompatibility complex social signalling in women Proc. R. Soc. B 285:20172714.

*** 本文描述了主要免疫遺傳學复合體（MHC）與人類氣味偏好之間的關聯，辯論了以往的發現，並分析了腋下刮鬍和其他因素如何影響身體氣味的感知及其與MHC的聯繫，重新分析過去的數據表明，氣味的愉悅度和強度可以與MHC的不相似性互動，從而影響通過氣味的社會信號，暗示MHC、氣味感知和社會溝通之間存在復雜的關係。

Wu K, Chen C, Moyzis RK, Nuno M, Yu Z and Greenberger E (2018) More than skin deep: Major histocompatibility complex (MHC)-based attraction among Asian American speed-daters Evolution and Human Behavior. 39(4):447-456.

*** 在速配約會的情境中，婦女會根據主要免疫相容性复合（MHC）通過SNP分析來偏好某些伴侶，而不同的SNP會根據與主要HLA基因的距離而有不同的吸引力，而MHC基準與自我報告的個性特徵在預測兩性的第二次約會邀請時有相當的影響力。

Henningsson S, Hovey D, Vass K, Walum H, Sandnabba K, Santtila P, Jern P, Westberg L (2017) A missense polymorphism in the putative pheromone receptor gene VN1R1 is associated with sociosexual behavior Transl Psychiatry. 7(4):e1102.

*** 研究發現，VN1R1基因的多型性與女性的社會性行為有顯著關聯，尤其是與一夜情有關，這表明化學信號可能在調節人類社會互動中起到作用，VN1R1基因與人類鼻腔型一受體1（費洛蒙作用系統的一部分）有關。

Burger D、Meuwly C、Marti E、Sieme H、Oberthür M、Janda J、Meinecke-Tillmann S 及 Wedekind C（2017） 「母馬（Equus caballus）在滯熱期間的MHC相關偏好。」

這項研究調查了主要組織相容性複合體（MHC）基因和睪酮水平對母馬交配偏好的影響，發現母馬在其黃體期（非繁殖期）對MHC不相似的種馬表現出更高的興趣，但其他與MHC無關的雄性特徵，可能如身體屬性或行為線索，可能在其發情期（繁殖期）中超越MHC的影響，情境和週期階段在伴侶選擇中起著關鍵作用。

Sherlock JM, Sidari MJ, Harris EA, Barlow FK, Zietsch BP (2016) Testing the mate-choice hypothesis of the female orgasm: disentangling traits and behaviours Socioaffect Neurosci Psychol. 6:31562.

*** 研究顯示，女性在與不同的伴侶交往時，對於高頻率的高潮反應會更加歡迎，因為這些伴侶會被認為更有幽默感、更有創造力、更溫暖、更忠誠，並且會更加關注伴侶的性滿足，部分支持女性高潮作為選擇高品質伴侶的機制的假設。

Kromer J, Hummel T, Pietrowski D, Giani AS, Sauter J, Ehninger G, Schmidt AH, Croy I (2016) Influence of HLA on human partnership and sexual satisfaction Sci Rep. 6:32550.

*** 研究表明，在人類中，HLA（免疫系統的一個遺傳成分）的不相似性可能通過檢測與基因相容性相關的嗅覺細胞來影響配偶選擇和性滿足，類似於其他動物中觀察到的機制，促進子代的基因多樣性和對病原體的抵抗力。

Wlodarski R. and Dunbar RIM. (2015) What's in a Kiss? The Effect of Romantic Kissing on Mate Desirability Evol Psychol. 12(1): 178–199.

*** 本研究發現，被認為的浪漫親吻能力會影響對潛在伴侶的評價，尤其是在非正式性行為情境中，對女性的吸引力會更高；而在某些交配情境中，視覺暗示可以抵銷親吻相關資訊的影響，但有吸引力的視覺資訊對男性的影響似乎比對女性更加強烈，這暗示親吻在傳達伴侶品質和吸引力時，與其他暗示一樣，扮演著多面向的角色。

Burger D、Dolivo G、Marti E、Sieme H 和 Wedekind C（2015） 「馬（Equus caballus）的雌性主要組織相容性复合體型對其雄性的睾丸素水平和精子數量有影響」

本研究旨在探討主要組織相容性复合物（MHC）如何影響馬的繁殖策略。研究發現，當雄馬暴露於MHC類型不同的母馬時，其睪酮水平較高，每次射精的精子數量也較多，與暴露於MHC類型相似的母馬相比，這顯示MHC相關的信號確實可以影響雄馬的睪酮生成和精液特性，從而影響繁殖策略。

Christakis NA and Fowler JH (2014) 友谊与自然选择 PNAS. 11:10796–10801.

*** 研究表明，人類往往會與擁有相似基因型的人形成友誼，相當於第四代表親，在整個基因組中，雖然有些基因型是正相關的（同質性），但也有些是負相關的（異質性），其中某些基因集與嗅覺和免疫系統有關，可能會影響友誼的形成，提出朋友可能充當“功能親屬”，並暗示同質性基因型可能提供協同適應優勢，這些優勢可能影響了近代人類的進化。

Laurent R and Chaix R (2012) MHC-dependent mate choice in humans: Why genomic patterns from the HapMap European American dataset support the hypothesis BioEssays. 34(4):267-71.

*** This study supports the hypothesis that humans tend to choose mates with different MHC (Major Histocompatibility Complex) genes than their own, which may promote genetic diversity and immune system strength in offspring.

Lie HC, Simmons LH and Rhodes G (2010) Genetic dissimilarity, genetic diversity, and mate preferences in humans Evolution and Human Behavior 31:8–58.

*** 本研究探討了遺傳因素，特別是主要組織相容性複合體（MHC），對人類伴侶偏好的影響，發現男性在短期和長期交配情境中傾向於偏好MHC不相似的女性，而遺傳多樣性影響了不同交配情境中男性和女性的伴侶偏好，從而支持MHC在人類伴侶選擇中的重要角色，並暗示這些偏好可能有助於增強後代的遺傳多樣性。

Lie HC, Rhodes G and Simmons LH (2010) Is genetic diversity associated with mating success in humans? Animal Behaviour. 79, 4:903-909

*** 這項研究表明，在人類中，特別是女性，基因多樣性，尤其是在主要組織相容性複合體 (MHC) 內，與更高的交配成功率相關，這是通過性伴侶的數量來衡量的，支持了基因因素，可能與免疫系統功能相關，對人類交配成功率起著作用的概念，而在男性中則未發現顯著的關聯。

Chaix R, Cao C, and Donnelly P (2008) Is Mate Choice in Humans MHC-Dependent? PLOS Genetics, 4 (9)

*** 研究表明，儘管歐洲裔美國人群可能偏好與MHC不相似的伴侶，以選擇多樣化的後代免疫系統，但在所檢查的非洲人群中並未發現這種模式，這表明MHC對伴侶選擇的影響可能是情境依賴的，並可能受到各種因素的影響。

Schwensow N, Fietz J, Dausmann K, Sommer S (2008) MHC-associated mating strategies and the importance of overall genetic diversity in an obligate pair-living primate Evol Ecol. (22) 617-636

*** 這項研究探討了肥尾矮狐猴的配偶選擇，發現雌性偏好具有較高MHC基因多樣性和較低MHC重疊的雄性作為社會和基因父親，而整體基因雜合性和親緣關係對配偶選擇並沒有顯著影響；額外配對可能發生以減輕基因不相容性，突顯了在靈長類動物配偶策略中「良好基因作為雜合性」和「不配對交配」假說之間的複雜相互作用。

Wedekind C (2007) The Major Histocompatibility Complex and Perfumers' Descriptions of Human Body Odors Evolutionary Psychology.5(2): 330-343

*** 這項實驗揭示了主要組織相容性複合體 (MHC) 與人類體味的口頭描述之間的聯繫，顯示專業香水師在某種程度上能夠表達與 MHC 相關的體味成分，強調了 MHC 對人類嗅覺感知的影響，以及在配偶選擇中的可能性，考慮到 MHC 在免疫系統功能中的關鍵角色以及之前提到的對體味和配偶選擇的影響。

Garver-Apgar CE, Gangestad SW, Thornill R, Miller RD and Olp JJ (2006) Major histocompatibility complex alleles, sexual responsivity, and unfaithfulness in romantic couples Psychol Sci, 17(10): 830-835.

*** In the context of romantic couples, the study finds that as the proportion of shared Major Histocompatibility Complex (MHC) alleles increases, women's sexual responsivity to their partners decreases, while their number of extra-pair sexual partners and attraction to men other than their primary partners, especially during their fertile phase, increases, suggesting that MHC-dissimilarity might play a role in sexual attraction and partner selection, potentially driven by underlying genetic and reproductive strategies to enhance offspring immunocompetence by maintaining genetic diversity.

Roberts CE, Gosling LM, Carter V and Petrie M (2006) MHC-correlated odour preferences in humans and the use of oral contraceptives Proc. R. Soc. B 275, 2715–2722

*** This study investigates how oral contraceptives influence women's preferences for certain male odours, which are known to be influenced by genes in the Major Histocompatibility Complex (MHC). Previous research has indicated that female preference for MHC-dissimilar male odour may function to increase offspring heterozygosity or minimize inbreeding. Interestingly, it has also been suggested that women using oral contraceptives tend to prefer the scent of MHC-similar males, which runs contrary to this logic. The study utilized a longitudinal design to test women before and after initiating contraceptive pill use, comparing their preferences to a control group of non-pill users. While the study didn't find a significant difference in ratings between odours of MHC-dissimilar and MHC-similar men during the follicular cycle phase in general, it did unearth that single women tended to prefer the odour of MHC-similar men, whereas women in relationships leaned towards the odour of MHC-dissimilar men. This latter finding aligns.

Wedekind C (2006) The Intensity of Human Body Odors and the MHC: Should We Expect A Link? Evolutionary Psychology. 4:85-94

*** 這項研究辨識出幾個MHC（主要組織相容性複合體）基因與男性體味的強度和愉悅感之間的關係，發現至少擁有一個純合子MHC抗原的男性並不會散發出顯著更強烈的氣味，相較於雜合子，但其氣味在擁有不同MHC的女性眼中卻被感知為顯著更強烈。

Pause BM, Krauel K, Schrader C, Sojka B, Westphal E, Müller-Ruchholtz W, and Ferstl R. (2006) The human brain is a detector of chemosensorily transmitted HLA-class I-similarity in same- and opposite-sex relations Proc. R. Soc. B (2006) 273, 471–478 doi:10.1098/rspb.2005.3342

*** This study reveals that humans subconsciously detect and respond to body odors from individuals with similar HLA (Human Leucocyte Antigen) genetic markers, which might play a critical role in social interactions and mate selection by acting as subtle social signals within both same-sex and opposite-sex relationships.

Martins Y, Preti G, Crabtree CR, Runyan T, Vainius AA and Wysocki CJ (2005) Preference for Human Body Odors Is Influenced by Gender and Sexual Orientation Psychological Science, 16(9) 694-702

*** 這項研究表明，性取向和性別影響個體對人體氣味的偏好，異性戀和同性戀的男性和女性在面對來自不同性別和性取向組合的個體的體味時，展示出明顯的二選一偏好，這暗示著體味可能在性伴侶和社交伴侶的選擇中扮演一個因素。

Horton R, Wilming L, Rand V, Lovering RC, Bruford EA, Khodiyar VK, Lush MJ, Povey S, Talbot CC Jr, Wright MW, Wain HM, Trowsdale J, Ziegler A, Beck S (2004) Gene map of the extended human MHC. Nat Rev Genet 5(12):889-899

*** This study characterises an integrated gene map of the extended human MHC, a pivotal region in the vertebrate genome for infection and autoimmunity due to its essential role in both adaptive and innate immunity, highlighting its significant content related to paralogy, polymorphism, immune functionality, and its association with various diseases, providing a comprehensive view stemming from the assimilation of data from several large-scale studies.

Jacob S, McClitock MK, Zelano B and Ober C (2002) Paternally inherited HLA alleles are associated with woman's choice of male odour Nature Genetics, 30:175

*** This study demonstrates that women can discern differences in male body odor influenced by a single variance in HLA (human MHC) allele, with a woman's capacity to distinguish and prefer certain odors notably being based on HLA alleles inherited from her father and not her mother; this suggests that paternally inherited HLA-associated odors influence odor preference and might function as social cues, even despite a milieu of potential odors derived from a variety of genetic and cultural factors.

Milinski M and Wedekind C (2001) Evidence for MHC-correlated perfume preference in humans Behavioural Ecology 12(2):140-149

*** 這項研究探討了主要組織相容性複合體（MHC）基因型與人類香水偏好之間的關係。他們發現個體的MHC（特別是HLA-A、-B、-DR）與其對各種香氣的個人使用評價之間存在顯著的相關性，某些HLA類型（例如，HLA-A2）顯示出一致的香氣偏好。然而，當談到對伴侶香氣的偏好時，相關性則大多不顯著，這與假設一致，即個體選擇香水用於個人使用，可能是為了增強或放大自身的免疫基因揭示的體味，而不是為了改變伴侶的香氣感知。

Wedekind C, Seebeck T, Bettens F And Paepke AJ (1995) MHC-dependent mate preferences in humans. Proc.R.Soc.Lond.B. 260:245-249.

*** 這項研究發現，人們傾向於偏好擁有不同免疫系統基因的潛在伴侶的體味，但這種偏好在使用避孕藥的女性中會有所改變。

* demonstrated increased rates of recurrent spontaneous abortion among Hutterite couples matching for HLA-DQA1 alleles

Ober., Steck., Ven., Billstrand., Messer., Kwak., Beaman., Beer (1993) MHC class II compatibility in aborted fetuses and term infants of couples with recurrent spontaneous abortion Journal of Reproductive Immunology, Volume 25, Issue 3, December 1993, Pages 195-207

* results of a 10 year study that demonstrated increased fetal loss rates among Hutterite couples matching for HLA antigens.

Ober C, Hyslop T, Elias S, Weitkamp LR, Hauck WW (1998) Human leukocyte antigen matching and fetal loss: results of a 10 year prospective study Human Reproduction, Volume 13, Issue 1, Jan 1998, Pages 33–38

* describe the relationship between HLA similarity at individual alleles, as well as HLA haplotypes as indicators for increased risk of fetal loss among Hutterite couples.

Ober C (1999) Studies of HLA, fertility and mate choice in a human isolate Hum Reprod Update 5(2):103-107.

* identified differential association of alleles in HLA-A, HLA-B, HLA-C, and HLA-DRB1 when compared with reported world RSA patients.

Shankarkumar U, Pawar A, Gaonkar P, Parasannavar D, Salvi V, and Ghosh K (2008) HLA allele associations in idiopathic recurrent spontaneous abortion patients from India J Hum Reprod Sci. 2008 Jan;1(1):19-24.

* found an increased frequency of homozygosity for HLA-E*0101 in Egyptian women with RM

Mosaad YM, Abdel-Dayem Y, El-Deek BS and El-Sherbini SM (2011) Association between HLA-E *0101 homozygosity and recurrent miscarriage in Egyptian women Scand J Immunol. 2011 Aug;74(2):205-9.

* performed a meta-analyses of 41 studies and showed that HLA-B sharing and HLA-DR sharing were both associated with the occurrence of RM.

Meuleman T, Lashley LE, Dekkers OM, van Lith J, Claas FH and Bloemenkamp KW (2015) HLA associations and HLA sharing in recurrent miscarriage: A systematic review and meta-analysis Hum Immunol. May;76(5):362-73.

* proposed a genetic-risk calculation that could predict recurrent miscarriage (RM) based on analysing HLA haplotypes from couples either with histories of successful pregnancies or RM.

Mora-Sánchez A, Aguilar-Salvador D, Nowak I (2019) Towards a gamete matching platform: using immunogenetics and artificial intelligence to predict recurrent miscarriage NPJ Digit Med Mar 7;2:12.

* found that sperm viability was associated with partners' HLA dissimilarity, indicating that cervical mucus may selectively facilitate later gamete fusion between immunogenetically compatible partners.

Jokiniemi A, Magris M, Ritari J, Kuusipalo L, Lundgren T, Partanen J and Kekäläinen J (2020) Post-copulatory genetic matchmaking: HLA-dependent effects of cervical mucus on human sperm function Proc Biol Sci., doi: 10.1098/rspb.2020.1682.

* demonstrated that HLA eplet dissimilarity positively affects sperm viability in follicular fluid, providing evidence that cryptic female choice operates at the gamete level to favour immunologically compatible partners.

Magris M, Jokiniemi A, Kuusipalo L, Ritari J, Koskela S, Partanen J and Kekäläinen J (2021) Structural dissimilarity of partners' immune genes increases sperm viability in women's reproductive tract Journal of Evolutionary Biology, 34(7):1125-1134.

* demonstrated that soluble HLA-G secreted by embryos into IVF culture medium correlates with implantation success and pregnancy rates, confirming the importance of HLA molecules in early pregnancy.

Lédée N, Petitbarat M, Chevrier L, et al. (2022) The impact of soluble HLA-G in IVF/ICSI embryo culture medium on implantation success Front Immunol. 13:982518.

* found that over 70% of couples with recurrent miscarriages or IVF failures showed high HLA-DQA1 allele similarity, supporting the role of HLA compatibility in reproductive outcomes.

Pafilis I, Michou V, Tsilivakos V, et al. (2023) Human Leukocyte Antigen Alleles Compatibility and Immunophenotypic Profile Associations in Infertile Couples Int J Mol Sci. 24(6):5350.

* found that primary infertile couples (N=609) share more HLA alleles than expected by chance, suggesting HLA similarity may contribute to unexplained infertility and IVF failure.

Kolańska K, Grześ S, Łukaszuk K, et al. (2025) Primary infertile couples share more HLA alleles than expected by chance European Journal of Obstetrics & Gynecology and Reproductive Biology.

* Landmark study demonstrating that house mice prefer mates with dissimilar MHC genotypes. Mice detect MHC differences through urine odor cues, and MHC-disassortative mating preferences result in a 27% deficit of MHC homozygous offspring in wild populations.

Penn DJ and Potts WK (1999) The evolution of mating preferences and major histocompatibility complex genes The American Naturalist, 153(2):145-164.

* Demonstrated that female three-spined sticklebacks (Gasterosteus aculeatus) use odor-based MHC detection to select mates that will produce offspring with optimal MHC diversity, equipping them with maximal resistance to pathogens and parasites.

Milinski M, Griffiths S, Wegner KM, Reusch TBH, Haas-Assenbaum A, Boehm T (2005) Mate choice decisions of stickleback females predictably modified by MHC peptide ligands PNAS, 102(12):4414-4418.

* Found that both male and female sticklebacks with intermediate MHC class IIB diversity had the highest lifetime reproductive success, demonstrating that optimal (not maximal) MHC diversity maximizes fitness.

Kalbe M, Eizaguirre C, Dankert I, Reusch TBH, Sommerfeld RD, Wegner KM, Milinski M (2009) Lifetime reproductive success is maximized with optimal major histocompatibility complex diversity Proceedings of the Royal Society B, 276(1658):925-934.

* In the grey partridge, females preferentially paired with males having more dissimilar MHC, with lower numbers of shared amino acid variants. This supports the 'inbreeding avoidance' and 'complementary genes' hypotheses in a strictly monogamous bird species.

Løvlie H, Gillingham MAF, Worley K, Pizzari T, Richardson DS (2017) Mate choice for major histocompatibility complex complementarity in a strictly monogamous bird, the grey partridge (Perdix perdix) Frontiers in Zoology, 14:9.

* Wild-spawning Atlantic salmon that chose their own mates produced offspring with 4× lower parasite loads than artificially crossed salmon, despite similar MHC diversity—demonstrating that MHC-mediated mate choice directly increases offspring fitness.

Consuegra S, de Leaniz CG (2008) MHC-mediated mate choice increases parasite resistance in salmon Proceedings of the Royal Society B, 275(1641):1397-1403.

* Discovered that MHC peptides trigger olfactory imprinting in zebrafish during a critical developmental period, creating persistent kin recognition preferences—demonstrating a fundamental mechanism by which vertebrates learn to recognize MHC-similar individuals.

Gerlach G, Hodgins-Davis A, Avolio C, Schunter C (2013) Olfactory imprinting is triggered by MHC peptide ligands Scientific Reports, 3:2800.

* Found non-random mating based on MHC in Chinook salmon—females directed more aggression toward MHC-similar males than MHC-dissimilar males, providing behavioral evidence for active mate discrimination based on immune genes.

Neff BD, Garner SR, Heath JW, Heath DD (2008) The MHC and non-random mating in a captive population of Chinook salmon Heredity, 101:175-185.

* In a study of 191 mares, females were significantly more likely to become pregnant when exposed to MHC-dissimilar stallions (p=0.019), demonstrating "cryptic female choice" influenced by MHC social signalling in horses.

Burger D, Meuwly C, Marti E, Sieme H, Oberthür M, Janda J, Meinecke-Tillmann S, Wedekind C (2017) MHC-linked social signalling affects female fertility in horses Proceedings of the Royal Society B, 284:20161998.

* Found that stallions exposed to MHC-dissimilar mares exhibited 19.2%% higher testosterone levels and produced more sperm per ejaculate, demonstrating that MHC-linked signals directly affect male reproductive strategies.

Burger D, Dolivo G, Marti E, Sieme H, Wedekind C (2015) 「馬（Equus caballus）的雌性主要組織相容性复合體型對其雄性的睾丸素水平和精子數量有影響」 Proceedings of the Royal Society B, 282:20150407.

* Meta-analysis of 58 effect sizes from 30 studies across seven primate species found a significant trend favouring MHC-diverse mates. The analysis confirms MHC-based mate preferences operate across the primate lineage.

Winternitz J, Abbate JL, Huchard E, Havlíček J, Garamszegi LZ (2017) Patterns of MHC-dependent mate selection in humans and nonhuman primates: a meta-analysis Molecular Ecology, 26(2):668-688.

* In wild grey mouse lemurs from Madagascar, found evidence for post-copulatory mate choice associated with MHC constitution—fathers had higher numbers of MHC supertypes different from mothers than randomly assigned males.

Schwensow N, Eberle M, Sommer S (2008) Compatibility counts: MHC-associated mate choice in a wild promiscuous primate Proceedings of the Royal Society B, 275(1634):555-564.

* Building on Burger's work, found that stallion semen quality depends on MHC matching to teaser mares—semen collected in the presence of MHC-dissimilar mares had higher sperm concentration and better motility.

Jeannerat E, Marti E, Berney C, Janett F, Bollwein H, Sieme H, Burger D, Wedekind C (2018) Stallion semen quality depends on major histocompatibility complex matching to teaser mare Molecular Ecology, 27(4):1025-1035.

* First evidence for MHC-based mate choice in reptiles: female Chinese alligators preferentially mate with MHC-heterozygous males that are genetically compatible, extending MHC mate choice to crocodilians.

Wang H, Shen FJ, Min MS, Wu XB, Yue BS, Yan P (2018) MHC class I diversity predicts non-random mating in Chinese alligators (Alligator sinensis) Heredity, 122:809-818.

* Found evidence for MHC-disassortative mating in the tuatara, a "living fossil" reptile—demonstrating that MHC-based mate choice has been conserved since the Triassic period (>200 million years ago).

Miller HC, Moore JA, Nelson NJ, Daugherty CH (2009) Influence of major histocompatibility complex genotype on mating success in a free-ranging reptile population Proceedings of the Royal Society B, 276(1662):1695-1704.

* In Seychelles warblers, when females' social mates were MHC-similar, they were more likely to engage in extra-pair copulations, with extra-pair males being significantly more MHC-dissimilar than social mates.

Richardson DS, Komdeur J, Burke T, von Schantz T (2005) MHC-based patterns of social and extra-pair mate choice in the Seychelles warbler Proceedings of the Royal Society B, 272(1564):759-767.

* Demonstrated that MHC-based mate choice in sticklebacks can drive ecological speciation, showing that immunogenes not only influence individual fitness but can shape evolutionary divergence between populations.

Eizaguirre C, Lenz TL, Kalbe M, Milinski M (2018) Mate choice in sticklebacks reveals that immunogenes can drive ecological speciation Behavioral Ecology, 28(4):953-961.

* Found in ring-necked pheasants that male spur length (a sexually selected ornament) was correlated with MHC genotype, demonstrating that sexual ornaments can signal genetic quality at immune genes.

von Schantz T, Wittzell H, Goransson G, Grahn M, Persson K (1996) MHC genotype and male ornamentation: genetic evidence for the Hamilton-Zuk model Proceedings of the Royal Society B, 263(1368):265-271.

* Quantitative review of MHC-based mating preferences across 50+ vertebrate species found significant effects for both MHC diversity and dissimilarity preferences, confirming MHC-mediated mate choice as a widespread vertebrate phenomenon.

Kamiya T, O'Dwyer K, Westerdahl H, Senior A, Nakagawa S (2014) A quantitative review of MHC-based mating preference: the role of diversity and dissimilarity Molecular Ecology, 23(21):5151-5163.

* Demonstrated MHC-based mate choice in sand lizards—females preferred MHC-dissimilar males, showing that reptiles can recognize both their own genotype and that of potential partners through scent cues.

Olsson M, Madsen T, Nordby J, Wapstra E, Ujvari B, Wittsell H (2003) Major histocompatibility complex and mate choice in sand lizards Proceedings of the Royal Society B, 270(Suppl 2):S254-S256.

* Found that frog tadpoles use MHC-based self-referent cues to preferentially associate with MHC-similar kin—demonstrating that MHC recognition operates in amphibians from early developmental stages.

Villinger J, Waldman B (2008) Self-referent MHC type matching in frog tadpoles Proceedings of the Royal Society B, 275(1640):1225-1230.

* Found that MHC-divergent male tiger salamanders had reduced mating success compared to males with intermediate MHC divergence—supporting the optimal MHC diversity hypothesis in amphibians.

Bos DH, Williams RN, Gopurenko D, Bulut Z, DeWoody JA (2009) Condition-dependent mate choice and a reproductive disadvantage for MHC-divergent male tiger salamanders Molecular Ecology, 18(15):3307-3315.

* Comprehensive review of MHC odor signaling mechanisms across vertebrates—synthesizes decades of research on how fish, mice, horses, and humans detect MHC differences through olfactory cues.

Milinski M (2022) A Review of Suggested Mechanisms of MHC Odor Signaling Biology, 11(8):1187.