Харилцааны нийцлийн шинжлэх ухаан

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.

Дандин-Руллан C, Лорант R, Даллара I, Тупанс B болон Шайкс R (2019) Хүний МHC-ийн ялгаатай хослолын геномын нотолгоо

Энэхүү судалгаа нь хүний ​​гол түүхийн нийцлийн комплекс (MHC) болон хамтрагчийн сонголтод хамааралтай бөгөөд Хойд Европын хосууд MHC-ний ялгаатай байдлыг харуулах хандлагатай байгааг харуулж байна. Энэ нь хамтрагчийн сонголтод биологийн нөлөө байж болохыг илтгэж байгаа ч Израиль зэрэг бусад бүс нутагт нийгмийн болон/эсвэл соёлын хүчин зүйлс ийм биологийн үндэслэлтэй гэрлэх хандлагыг давж гарах эсвэл бүр бүрхэг болгох боломжтой юм.

Birnbaum GE, Zholtack K, Mizrahi M & Ein-Dor T (2019) Гашуун эм: Амны контрацептивыг зогсоосноор орлуулах хамтрагчдын таталцлыг нэмэгдүүлдэг Evolutionary Psychological Science volume 5, pages 276–285

*** Гормоны зохицуулах эмүүд эмэгтэйчүүдийн хосын сонголтыг илүү халамжтай эрчүүд рүү өөрчилдөг боловч эдгээрийг хэрэглэхээ больсноор эдгээр сонголтууд буцаж эргэж, генетикийн хувьд илүү нийцтэй эрчүүд рүү татагдахыг нэмэгдүүлдэг. Ялангуяа өндөр үржил шимтэй үеийн үед энэ нь өөр хосуудыг хайхыг урамшуулж болзошгүй.

Wedekind C (2019) Сайхан үнэрийн тааламжтай байдал болон хүчтэй байдлын хоорондын урьдчилан таамагласан харилцан үйлчлэл нь эмэгтэйчүүдийн хувьд гол тэжээллэг нийцлийн комплексийн нийгмийн дохио зангаа илтгэж байна. Proc. R. Soc. B 285:20172714.

*** Энэхүү судалгаа нь Хүний Төвд Тохиромжтой Комплекс (MHC) болон хүний үнэрийн сонголтын хоорондын холбоог тайлбарлаж, өмнөх судалгааны үр дүнгүүдийг хэлэлцэж, гараар хумслах болон бусад хүчин зүйлүүд нь биеийн үнэрийн ойлголтод хэрхэн нөлөөлж, MHC-тай хэрхэн холбогдож байгааг шинжилж байна. Өмнөх өгөгдлийг дахин шинжилснээр үнэрийн тааламжтай байдал болон хүчтэй байдал нь MHC-ийн ялгаатай байдлыг нийгмийн дохиололд үнэрээр нөлөөлөхөд хэрхэн харилцан үйлчилж болохыг илрүүлж, MHC, үнэрийн ойлголт, нийгмийн харилцааны хооронд нарийн төвөгтэй холбоо байгааг илтгэж байна.

Wu K, Chen C, Moyzis RK, Nuno M, Yu Z and Greenberger E (2018) Харилцаа холбоо нь арьснаас гүнзгий: Азийн Америкийн хурдан болзогчдын дунд гол гистосумын нийцлийн комплекс (MHC)-д суурилсан таталцал Evolution and Human Behavior. 39(4):447-456.

*** Хурдан болзоо (speed-dating) контекстод эмэгтэйчүүд, эрэгтэйчүүд биш, Махжийн түүхийн нийцлийн комплекс (MHC) нийцлийн үндсэн дээр түншүүдийг сонгох сонирхолтой байв. SNP шинжилгээний үр дүнд янз бүрийн SNP-ууд HLA генүүдийн ойртох байдалд үндэслэн таталцлын чиглэлийг нөлөөлж байсан бөгөөд MHC-д суурилсан индексүүд нь хоёр хүйсийн хувьд хоёр дахь болзоо санал болгосон таамаглалд өөрийгөө тайлагнасан зан чанартай адил нөлөө үзүүлж байв.

Henningsson S, Hovey D, Vass K, Walum H, Sandnabba K, Santtila P, Jern P, Westberg L (2017) VN1R1 генийн таамагласан феромон хүлээн авагчид missense полиморфизм нь нийгмийн бэлгийн зан үйлтэй холбоотой байна. Transl Psychiatry. 7(4):e1102.

*** Судалгаа нь хүний вомероназаль төрөл-1 хүлээн авагч 1 (феромоны нөлөөг дамжуулах системийн нэг хэсэг) VN1R1 генийн полиморфизм болон эмэгтэйчүүдийн нийгмийн бэлгийн зан үйл, ялангуяа нэг шөнийн харилцаатай холбоотойгоор чухал холбоос байгааг олж тогтоожээ. Энэ нь химийн дохиолол нь хүний нийгмийн харилцааг зохицуулахад үүрэг гүйцэтгэж болохыг санал болгож байна.

Бургер Д, Меуэли С, Марти Е, Сиеме Х, Обертюр М, Жанда Ж, Мейнеке-Тиллманн С, Ведекинд С. (2017) Диэстроз үеийн эм х馬 (Equus caballus)-ийн MHC-тай холбоотой предпочтения.

Энэ судалгаа нь Үндсэн Гистосумын Тогтолцоо (MHC) генүүд болон тестостероны түвшин эмэгтэй морьдын хослолын сонголтод хэрхэн нөлөөлдгийг судалсан бөгөөд эмэгтэй морьд MHC-тэй адилгүй эр морьдод ихээхэн сонирхолтой ханддаг болохыг олж мэдсэн. Энэ нь тэдний диэстрозын үе (бэлгийн үржилд орохгүй үе) үед ажиглагдсан. Гэвч бусад MHC-тэй холбоогүй эрэгтэй шинж чанарууд, магадгүй биеийн онцлог эсвэл зан төлөвийн дохио зэрэг нь эмэгтэй морьд MHC-ийн нөлөөг давж гарах боломжтой бөгөөд энэ нь тэдний эстрозын (бэлгийн үржил) үеийн хослолын сонголтод чухал үүрэгтэй байдаг.

Sherlock JM, Sidari MJ, Harris EA, Barlow FK, Zietsch BP (2016) Эмэгтэйчүүдийн оргасмын хос сонголтын гипотезийг турших: шинж чанар, зан үйлийг задлах Socioaffect Neurosci Psychol. 6:31562.

*** Судалгаанаас үзэхэд, эмэгтэйчүүд нь янз бүрийн түншүүдтэйгээ харилцахад өөр өөр давтамжийн оргасм хүртдэг бөгөөд өндөр оргасм үүсгэдэг түншүүдийг илүү инээдтэй, бүтээлч, дулаан, үнэнч, сайхан үнэртэй гэж үнэлдэг бөгөөд мөн түншийнхээ бэлгийн сэтгэл ханамжид илүү анхааралтай ханддаг. Энэ нь эмэгтэйчүүдийн оргасмын хос сонголтын гипотезийг тодорхой хэмжээгээр дэмжиж байгаа бөгөөд энэ нь өндөр чанартай түншүүдийг сонгох боломжит механизм гэж үздэг.

Kromer J, Hummel T, Pietrowski D, Giani AS, Sauter J, Ehninger G, Schmidt AH, Croy I (2016) HLA-ийн хүний түншлэл болон бэлгийн сэтгэл ханамжид үзүүлэх нөлөө Sci Rep. 6:32550.

*** The study indicates that in humans, HLA (a genetic component of the immune system) dissimilarity can influence mate selection and sexual satisfaction, potentially through the detection of olfactory cues related to genetic compatibility, similarly to mechanisms observed in other animals, promoting genetic diversity and resilience against pathogens in offspring.

Wlodarski R. and Dunbar RIM. (2015) Цалингийн утга юу вэ? Романтик үнсэлт нь хамтрагчийн хүсэлд хэрхэн нөлөөлдөг вэ? Evol Psychol. 12(1): 178–199.

*** This study found that perceived romantic kissing ability affects evaluations of potential mates, increasing desirability particularly for women in casual sex contexts, and while visual cues can override the impact of kissing-related information in certain mating scenarios, the influence of attractive visual information appears more potent for men than for women, alluding to the multifaceted role of kissing in conveying mate quality and desirability alongside other cues.

Бургер Д, Доливо Г, Марти Е, Сиеме Х, Ведекинд С. (2015) Эмэгтэй гол гистосумын нийцлийн комплексийн төрөл нь морь (Equus caballus)-ийн эрэгтэй тестостероны түвшин болон үрийн тоонд нөлөөлдөг.

This study examines how the Major Histocompatibility Complex (MHC) influences male reproductive strategies in horses. The research revealed that when stallions were exposed to females with dissimilar MHC types, they exhibited higher testosterone levels and produced a larger number of sperm per ejaculate compared to when they were exposed to MHC-similar mares, highlighting that MHC-linked signals can indeed impact male testosterone production and semen characteristics, thereby influencing reproductive approaches.

Christakis NA and Fowler JH (2014) Найрамдал ба байгалийн сонголт PNAS. 11:10796–10801.

*** The study demonstrates that humans tend to form friendships with individuals who have similar genotypes—akin to the level of fourth cousins—across the entire genome, and while certain genotypes are positively correlated (homophilic), others are negatively correlated (heterophilic) among friends, with particular gene sets related to olfactory and immune system playing a role in friendship formation, proposing that friends might act as "functional kin" and suggesting that homophilic genotypes may offer synergistic fitness benefits that have been influencing recent human evolution.

Laurent R and Chaix R (2012) Хүний MHC-д тулгуурласан хос сонголт: HapMap Европын Америкчуудын мэдээллийн сангаас гарсан геномын загварууд яагаад гипотезыг дэмжиж байгааг тайлбарлах нь 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) Генетикийн ялгаа, генетикийн олон янз байдал, болон хүний хослолын сонголтууд Evolution and Human Behavior 31:8–58.

*** This study explores the influence of genetic factors, particularly the major histocompatibility complex (MHC), on mate preferences in humans, discovering that males tend to prefer MHC-dissimilar females in both short-term and long-term mating contexts, while genetic diversity impacts both male and female partner preferences across different mating contexts, thus supporting a significant role for the MHC in human mate selection and suggesting that these preferences might function to enhance genetic diversity in offspring.

Lie HC, Rhodes G and Simmons LH (2010) Генетикийн олон янз байдал нь хүний ​​хослолын амжилттай холбоотой юу?? Animal Behaviour. 79, 4:903-909

*** This study indicates that in humans, particularly females, genetic diversity, especially within the major histocompatibility complex (MHC), is associated with greater mating success, measured by the number of sexual partners, supporting the concept that genetic factors, potentially related to immune system functioning, play a role in human mating success, while no significant association was found in males.

Chaix R, Cao C, and Donnelly P (2008) Хүний хамтрагчийн сонголт MHC-д хамааралтай юу?? PLOS Genetics, 4 (9)

*** The study indicates that while European American populations may prefer MHC-dissimilar mates, selecting for diversity in offspring's immune systems, such a pattern is not found in the examined African population, suggesting MHC's influence on mate choice can be context-dependent and potentially shaped by various factors.

Schwensow N, Fietz J, Dausmann K, Sommer S (2008) MHC-тай холбоотой хослолын стратегиуд ба зайлшгүй хослон амьдрах приматад генетикийн олон янз байдлын чухал ач холбогдол Evol Ecol. (22) 617-636

*** The study explores mate choice in the fat-tailed dwarf lemur, finding females prefer males with greater MHC-genetic diversity and lower MHC overlap as both social and genetic fathers, while overall genetic heterozygosity and relatedness do not significantly influence mate selection; extra-pair matings might occur to mitigate genetic incompatibility, highlighting a complex interplay of "good-genes-as-heterozygosity" and "disassortative mating" hypotheses in primate mating strategies.

Wedekind C (2007) Гол Тулгарлын Комплекс ба Үнэрчдийн Хүний Биеийн Үнэрийн Тодорхойлолтууд Evolutionary Psychology.5(2): 330-343

*** This experiment reveals a connection between the Major Histocompatibility Complex (MHC) and the verbal descriptions of human body odors, demonstrating that professional perfumers can, to some extent, articulate MHC-correlated body odor components, underscoring the MHC's influence on human olfactory perception and possibly, mate selection, given the MHC's critical role in immune system functioning and previously noted impact on body odor and mate choice.

Garver-Apgar CE, Gangestad SW, Thornill R, Miller RD and Olp JJ (2006) Гол гистосумын нийцлийн комплексийн аллель, бэлгийн хариу үйлдэл, болон хайртай хосуудын үнэнч бус байдал 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) Хүний MHC-тай холбогдсон үнэрийн сонголт ба амны бэлгийн хяналтын хэрэгслийн хэрэглээ 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) Хүний биеийн үнэрийн хүчтэй байдал ба MHC: Холбоо байхыг хүлээх ёстой юу?? Evolutionary Psychology. 4:85-94

*** This research discerns relationships between a few MHC (Major Histocompatibility Complex) genes and the intensity and pleasantness of men's body odors, discovering that men with at least one homozygous MHC antigen don't emit notably more intense odors than heterozygotes, yet their scents are perceived as significantly stronger by women with dissimilar MHC.

Pause BM, Krauel K, Schrader C, Sojka B, Westphal E, Müller-Ruchholtz W, and Ferstl R. (2006) Хүний тархи нь ижил болон эсрэг хүйсийн харилцаанд химийн мэдрэгчээр дамжуулсан HLA-классын I-төрлийн ижил байдлыг илрүүлэгч юм. Proc. R. Soc. B (2006) 273, 471–478 doi:10.1098/rspb.2005.3342

*** Энэхүү судалгаа нь хүмүүс HLA (Хүний лейкоцитын антиген) генетикийн маркертай ижил хүмүүсийн биеийн үнэрийг ухамсаргүйгээр илрүүлж, хариу үйлдэл үзүүлдэгийг илрүүлжээ. Энэ нь нийгмийн харилцаа, хос сонголтод чухал үүрэг гүйцэтгэж, ижил хүйсийн болон эсрэг хүйсийн харилцаанд нарийн нийгмийн дохио болгон үйлчилж болох юм.

Martins Y, Preti G, Crabtree CR, Runyan T, Vainius AA and Wysocki CJ (2005) Хүний биеийн үнэрт дурлах нь хүйс болон бэлгийн чиг баримтлалтай холбоотойгоор нөлөөлдөг. Psychological Science, 16(9) 694-702

*** This study demonstrates that sexual orientation and gender influence individuals' preferences for human body odors, with heterosexual and homosexual males and females showcasing distinct two-choice preferences when presented with body odors from individuals of different gender and sexual orientation combinations, implying that body odor can be a factor in sexual and social partner selection.

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) Хүний өргөтгөсөн генийн зураг 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) Хүний MHC-тай холбоотой үнэрийн сонголтын нотолгоо Behavioural Ecology 12(2):140-149

*** The study explores the relationship between Major Histocompatibility Complex (MHC) genotypes and perfume preferences in humans. They discovered a notable correlation between an individual's MHC (specifically HLA-A, -B, -DR) and their ratings of various scents for personal use, with certain HLA types (e.g., HLA-A2) showing consistent scent preferences. However, when it came to preferences for a partner's scent, the correlation was largely insignificant, aligning with the hypothesis that individuals select perfumes for personal use to possibly enhance or amplify their own immunogenetic revealing body odors, rather than to alter the perceived scents of their partners.

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

*** Энэ судалгаа нь хүмүүс өөрийнх нь дархлаа системийн генүүдээс ялгаатай потенциал хосуудын биеийн үнэрийг илүүд үздэг болохыг олж тогтоосон боловч, энэ илүүд үзэлт нь жирэмслэлтээс хамгаалах эм ууж буй эмэгтэйчүүдэд өөрчлөгддөг.

* 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) Египет эмэгтэйчүүдийн дахин давтагдсан зулбалттай холбоотой HLA-E *0101 гомозигот байдал 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) Эмэгтэй гол гистосумын нийцлийн комплексийн төрөл нь морь (Equus caballus)-ийн эрэгтэй тестостероны түвшин болон үрийн тоонд нөлөөлдөг. 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.