注释:

1. Ayala was not the only one to do this. See N. Takahata , “Allelic Genealogy and human evolution,” Mol Biol Evol 10 (1993): 2–22.

2. Briefly, HLA-DRB1 has six exons (the coding regions) interspersed by noncoding DNA, called introns.

3. Phylogenetics is the study of evolutionary relationships among organisms.These relationships are often represented as branching trees. Starting with the assumption that common descent is true, scientists compare the distribution of varying anatomical traits or DNA sequences that they are studying. Usingmathematical algorithms, they look for tree-branching patterns that minimize conflict, or represent the fewest changes over time, but that can explain the observed distribution of traits or DNA variation.

4. Francisco Ayala, “The myth of Eve: Molecular biology and human origins,” Science 270 (1995): 1930–1936.

5. H. A. Erlich et al., “HLA sequence polymorphism and the origin of humans,” Science 274 (1996): 1552–1554.

6. T. F. Bergström et al., “Recent origin of HLA-DRB1 alleles and implications for human evolution,” Nature Genetics 18 (1998): 237–242.

7. G. Doxiadis et al., “Reshuffling of ancient peptide binding motifs between HLA-DRB multigene family members: Old wine served in new skins,” Molecular Immunology 45 (2008): 2743–2751.

8. Ibid.

9. J. Klein, A. Sato, and N. Nikolaidis, “MHC, TSP, and the Origin of Species: From Immunogenetics to Evolutionary Genetics,” Annu. Rev. Genet. 41 (2007): 281–304.

10. Doxiadis, “Reshuffling of ancient peptide binding motifs.”

11. C.K. Raymond et al., “Ancient haplotypes of the HLA Class II region,” Genome Research 15 (2005): 1250–1257.

12. There is an illustration of HLA-DRB1 and its neighboring genes in C. K. Raymond et al., “Ancient haplotypes,” 1251.

13. G. Andersson, “Evolution of the human HLA-DR region,” Frontiers in Bioscience 3 (1998): d739–745.

14. V. Vincek, et al., “How Large Was the Founding Population of Darwin’s Finches?” Proc. R. Soc. London Ser. B 264 (1997): 111–118.

15. G. Doxiadis et al., “Extensive DRB region diversity in cynomolgus macaques: recombination as a driving force,” Immunogenetics 62 (2010): 137–147.

16. Ziqiang Li, Caroline J. Woo, Maria D. Iglesias-Ussel, et al.,“The generation of antibody diversity through hypermutation and class switch recombination,” Genes Dev. 18 (2004): 1–11.

17. Katja Kotsch and Rainer Blasczy, “Interlineage Recombinations as a Mechanism of The Noncoding Regions of HLA-DRB Uncover HLA Diversification,” J Immunol 165 (2000): 5664–5670.

18. Jenny von Salomé and Jyrki P Kukkonen, “Sequence features of HLA-DRB1 locus define putative basis for gene conversion and point mutations,” BMC Genomics 9 (2008): 228, accessed March 6, 2012, doi:10.1186/1471-2164-9-228.

19. P. W. Hedrick and T. Kim, “Genetics of Complex Polymorphisms: Parasites and Maintenance of the Major Histocompability Complex Variation,” in R. S. Singh and C. B. Crimbas, editors, Evolutionary Genetics: from Molecules to Morphology (New York: Cambridge University Press, 2000), 211–212.

20. E. A. Titus-Trachtenberg, et al., “Analysis of HLA Class 11 Haplotypes in the Cayapa Indians of Ecuador: A Novel DRB1 Allele Reveals Evidence for Convergent Evolution and Balancing Selection at Position 86,” Am. J. Hum. Genet. 55 (1994):160–167.

21. Hedrick and Kim, “Genetics of Complex Polymorphisms”; Gabriele Zangenberg, et al., “New HLA–DPB1 alleles generated by interallelic gene conversion detected by analysis of sperm,” Nature Genetics 10 (1995): 407–414, accessed March 6, 2012, doi:10.1038/ng0895-407.

22. A. Hobolth, O. F. Christensen, T. Mailund, M. H. Schierup, “Genomic Relationships and Speciation Times of Human, Chimpanzee, and Gorilla Inferred from a Coalescent Hidden Markov Model,” PLoS Genet 3 (2007): e7, accessed March 6, 2012,doi:10.1371/journal.pgen.0030007.