Identification of novel susceptibility loci for Guam neurodegenerative disease: Challenges of genome scans in genetic isolates

Human Molecular Genetics

Volumn 18, Issue 19, 2009, Pages 3725-3738

  Sieh, Weiva ^a,c   Choi, Yoonha ^b   Chapman, Nicola H ^a   Craig, Ulla Katrina ^d   Steinbart, Ellen J ^a,e   Rothstein, Joseph H ^a   Oyanagi, Kiyomitsu ^f   Garruto, Ralph M ^g   Bird, Thomas D ^a,e   Galasko, Douglas R ^h   Schellenberg, Gerard D ^a,e,i   Wijsman, Ellen M ^a,b  

^a UNIVERSITY OF WASHINGTON SCHOOL OF MEDICINE   (United States)

^b UNIVERSITY OF WASHINGTON   (United States)

^c STANFORD UNIVERSITY   (United States)

^d UNIVERSITY OF GUAM   (Guam)

^e VA PUGET SOUND HEALTH CARE SYSTEM   (United States)

^f TOKYO METROPOLITAN INSTITUTE OF MEDICAL SCIENCE   (Japan)

^g BINGHAMTON UNIVERSITY   (United States)

^h UNIVERSITY OF CALIFORNIA   (United States)

ⁱ UNIVERSITY OF PENNSYLVANIA   (United States)

Author keywords

[No Author keywords available]

Indexed keywords

ABORIGINE; ADULT; AGED; ARTICLE; CASE CONTROL STUDY; CHROMOSOME 12; CHROMOSOME 17; DEGENERATIVE DISEASE; FEMALE; GENETIC ASSOCIATION; GENETIC LINKAGE; GENETIC PREDISPOSITION; GENETICS; GUAM; HUMAN; MALE; MIDDLE AGED; PEDIGREE;

ADULT; AGED; CASE-CONTROL STUDIES; CHROMOSOMES, HUMAN, PAIR 12; CHROMOSOMES, HUMAN, PAIR 17; FEMALE; GENETIC PREDISPOSITION TO DISEASE; GENOME-WIDE ASSOCIATION STUDY; GUAM; HUMANS; LINKAGE (GENETICS); MALE; MIDDLE AGED; NEURODEGENERATIVE DISEASES; OCEANIC ANCESTRY GROUP; PEDIGREE; YOUNG ADULT;

EID: 70350764965     PISSN: 09646906     EISSN: 14602083     Source Type: Journal    
DOI: 10.1093/hmg/ddp300     Document Type: Article

References (95)

1. Hirano, A., Krooth, R.S., Lessell, S. and Kurland, L.T. (1961) Parkinsonism-dementia complex, an endemic disease on the island of Guam. 1. Clinical features. Brain, 84, 642-661.
2. Koerner, D.R. (1952) Amyotrophic lateral sclerosis on Guam-a clinical study and review of the literature. Ann. Intern. Med., 37, 1204-1220.
3. Kurland, L.T. and Mulder, D.W. (1954) Epidemiologic investigations of amyotrophic lateral sclerosis. 1. Preliminary report on geographic distribution, with special reference to the Mariana Islands, including clinical and pathologic observations. Neurology, 4, 438-448.
4. Kuzuhara, S. and Kokubo, Y. (2005) Atypical parkinsonism of Japan: amyotrophic lateral sclerosis-parkinsonism-dementia complex of the Kii peninsula of Japan (Muro disease): an update.Mov. Disord., 20, S108-S113.
5. Mimuro, M., Kokubo, Y. and Kuzuhara, S. (2007) Similar topographical distribution of neurofibrillary tangles in amyotrophic lateral sclerosis and parkinsonism-dementia complex in people living in the Kii peninsula of Japan suggests a single tauopathy. Acta Neuropathol. (Berl.), 113, 653-658.
6. Gajdusek, D.C. and Salazar, A.M. (1982) Amyotrophic lateral sclerosis and parkinsonian syndromes in high-incidence among the Auyu and Jakai people of west New-Guinea. Neurology, 32, 107-126.
7. Hirano, A., Malamud, N. and Kurland, L.T. (1961) Parkinsonism-dementia complex, an endemic disease on the island of Guam. 2. Pathological features. Brain, 84, 662-679.
8. Hirano, A., Malamud, N., Elizan, T.S. and Kurland, L.T. (1966) Amyotrophic lateral sclerosis and parkinsonism-dementia complex on Guam-further pathologic studies. Arch. Neurol., 15, 35-51.
9. Hof, P.R., Nimchinsky, E.A., Bueescherrer, V., Buee, L., Nasrallah, J., Hottinger, A.F., Purohit, D.P., Loerzel, A.J., Steele, J.C., Delacourte, A. et al. (1994) Amyotrophic-lateral-sclerosis parkinsonism-dementia complex of Guam-quantitative neuropathology, immunohistochemical analysis of neuronal vulnerability, and comparison with related neurodegenerative disorders. Acta Neuropathol. (Berl.), 88, 397-404.
10. Hof, P.R. and Perl, D.P. (2002) Neurofibrillary tangles in the primary motor cortex in Guamanian amyotrophic lateral sclerosis/ parkinsonism-dementia complex. Neurosci. Lett., 328, 294-298.
11. Hirano, A., Arumugas, N. and Zimmerma, H.M. (1967) Amyotrophic lateral sclerosis-a comparison of Guam and classical cases. Arch. Neurol., 16, 357-363.
12. Lessell, S., Torres, J., Kurland, L.T. and Hirano, A. (1962) Parkinsonism-dementia complex-epidemiological considerations in Chamorros of Mariana Islands and California. Arch. Neurol., 7, 377-385.
13. Reed, D.M. and Brody, J.A. (1975) Amyotrophic lateral sclerosis and parkinsonism-dementia on Guam, 1945-1972. 1. Descriptive epidemiology. Am. J. Epidemiol., 101, 287-301.
14. Plato, C.C., Garruto, R.M., Galasko, D., Craig, U.K., Plato, M., Gamst, A., Torres, J.M. and Wiederholt, W. (2003) Amyotrophic lateral sclerosis and parkinsonism-dementia complex of Guam: changing incidence rates during the past 60 years. Am. J. Epidemiol., 157, 149-157.
15. Garruto, R.M., Yanagihara, R. and Gajdusek, D.C. (1985) Disappearance of high-incidence amyotrophic lateral sclerosis and parkinsonism-dementia on Guam. Neurology, 35, 193-198.
16. Galasko, D., Salmon, D.P., Craig, U.K., Thal, L.J., Schellenberg, G. and Wiederholt, W. (2002) Clinical features and changing patterns of neurodegenerative disorders on Guam, 1997-2000. Neurology, 58, 90-97.
17. Gibbs, C.J. Jr and Gajdusek, D.C. (1972) Amyotrophic lateral sclerosis, Parkinson's disease, and the amyotrophic lateral sclerosis-Parkinsonism-dementia complex on Guam: a review and summary of attempts to demonstrate infection as the aetiology. J. Clin. Pathol. Suppl. (R Coll Pathol), 6, 132-140.
18. Spencer, P.S., Nunn, P.B., Hugon, J., Ludolph, A.C., Ross, S.M., Roy, D.N. and Robertson, R.C. (1987) Guam amyotrophic-lateral-sclerosis parkinsonism dementia linked to a plant excitant neurotoxin. Science, 237, 517-522.
19. Garruto, R.M., Fukatsu, R., Yanagihara, R., Gajdusek, D.C., Hook, G. and Fiori, C.E. (1984) Imaging of calcium and aluminum in neurofibrillary tangle-bearing neurons in parkinsonism-dementia of Guam. Proc. Natl Acad. Sci. USA, 81, 1875-1879.
20. Yanagihara, R., Garruto, R.M., Gajdusek, D.C., Tomita, A., Uchikawa, T., Konagaya, Y., Chen, K.M., Sobue, I., Plato, C.C. and Gibbs, C.J. (1984) Calcium and vitamin-D metabolism in Guamanian Chamorros with amyotrophic lateral sclerosis and parkinsonism-dementia. Ann. Neurol., 15, 42-48.
21. Baileywilson, J.E., Plato, C.C., Elston, R.C. and Garruto, R.M. (1993) Potential role of an additive genetic component in the cause of amyotrophic-lateral-sclerosis and parkinsonism-dementia in the Western Pacific. Am. J. Med. Genet., 45, 68-76.
22. Plato, C.C., Reed, D.M., Elizan, T.S. and Kurland, L.T. (1967) Amyotrophic lateral sclerosis/parkinsonism-dementia complex of Guam. 4. Familial and genetic investigations. Am. J. Hum. Genet., 19, 617-632.
23. Reed, D.M., Torres, J.M. and Brody, J.A. (1975) Amyotrophic lateral sclerosis and parkinsonism-dementia on Guam, 1945-1972. 2. Familial and genetic studies. Am. J. Epidemiol., 101, 302-310.
24. Plato, C.C., Galasko, D., Garruto, R.M., Plato, M., Gamst, A., Craig, U.K., Torres, J.M. and Wiederholt, W. (2002) ALS and PDC of Guam- forty-year follow-up. Neurology, 58, 765-773.
25. Plato, C.C., Cruz, M.T. and Kurland, L.T. (1969) Amyotrophic lateral sclerosis-parkinsonism dementia complex of Guam: further genetic investigations. Am. J. Hum. Genet., 21, 133-141.
26. Vilar, M., Lynch, D., Chan, C., Reiff, D., Garruto, R.M. and Lum, J.K. (2008) Peopling of the Marianas Islands: a mtDNA perspective. Am. J. Hum. Biol., 20, 237-237.
27. Lum, J.K. and Cann, R.L. (2000) mtDNA lineage analyses: origins and migrations of Micronesians and Polynesians. Am. J. Phys. Anthropol., 113, 151-168.
28. Rogers, R.F. (1995) Destiny's Landfall: A History of Guam. University of Hawaii Press, Honolulu.
29. Underwood, J.H. (1976) The native origins of the neo-Chamorros of the Mariana Islands. Micronesia, 12, 203-209.
30. Yanagihara, R.T., Garruto, R.M. and Gajdusek, D.C. (1983) Epidemiological surveillance of amyotrophic lateral sclerosis and parkinsonism-dementia in the Commonwealth of the Northern Mariana Islands. Ann. Neurol., 13, 79-86.
31. Bourgain, C. and Genin, E. (2005) Complex trait mapping in isolated populations: are specific statistical methods required? Eur. J. Hum. Genet., 13, 698-706.
32. Peltonen, L., Palotie, A. and Lange, K. (2000) Use of population isolates for mapping complex traits. Nat. Rev. Genet., 1, 182-190.
33. Service, S., DeYoung, J., Karayiorgou, M., Roos, J.L., Pretorious, H., Bedoya, G., Ospina, J., Ruiz-Linares, A., Macedo, A., Palha, J.A. et al. (2006) Magnitude and distribution of linkage disequilibrium in population isolates and implications for genome-wide association studies. Nat. Genet., 38, 556-560.
34. Venken, T. and Del-Favero, J. (2007) Chasing genes for mood disorders and schizophrenia in genetically isolated populations. Hum. Mutat., 28, 1156-1170.
35. Greenberg, D.A., Abreu, P. and Hodge, S.E. (1998) The power to detect linkage in complex disease by means of simple LOD-score analyses. Am. J. Hum. Genet., 63, 870-879.
36. Liu, F., Arias-Vasquez, A., Sleegers, K., Aulchenko, Y.S., Kayser, M., Sanchez-Juan, P., Feng, B.J., Bertoli-Avella, A.M., van Swieten, J., Axenovich, T.I. et al. (2007) A genomewide screen for late-onset Alzheimer disease in a genetically isolated Dutch population. Am. J. Hum. Genet., 81, 17-31.
37. Wijsman, E.M. and Amos, C.I. (1997) Genetic analysis of simulated oligogenic traits in nuclear and extended pedigrees: summary of GAW10 contributions. Genet. Epidemiol., 14, 719-735.
38. Chapman, N., Leutenegger, A.-L., Badzioch, M., Bogdan, M., Conlon, E., Daw, E., Gagnon, F., Li, N., Maia, J., Wijsman, E. et al. (2001) The importance of connections: joining components of the Hutterite pedigree. Genet. Epidemiol., 21, S230-S235.
39. Dyer, T., Blangero, J., Williams, J., Göring, H. and Mahaney, M. (2001) The effect of pedigree complexity on quantitative trait linkage analysis. Genet. Epidemiol., 21, S236-S243.
40. Leutenegger, A.L., Genin, E., Thompson, E.A. and Clerget-Darpoux, F. (2002) Impact of parental relationships in maximum LOD score affected sib-pair method. Genet. Epidemiol., 23, 413-425.
41. Genin, E. and Clerget Darpoux, F. (1996) Consanguinity and the sib-pair method: an approach using identity by descent between and within individuals. Am. J. Hum. Genet., 59, 1149-1162.
42. Devlin, B. and Roeder, K. (1999) Genomic control for association studies. Biometrics, 55, 997-1004.
43. Voight, B.F. and Pritchard, J.K. (2005) Confounding from cryptic relatedness in case-control association studies. PLoS Genet., 1, 302-311.
44. Chanock, S.J., Manolio, T., Boehnke, M., Boerwinkle, E., Hunter, D.J., Thomas, G., Hirschhorn, J.N., Abecasis, G., Altshuler, D., Bailey-Wilson, J.E. et al. (2007) Replicating genotype-phenotype associations. Nature, 447, 655-660.
45. Morris, H.R., Steele, J.C., Crook, R., Wavrant-De Vrieze, F., Onstead-Cardinale, L., Gwinn-Hardy, K., Wood, N.W., Farrer, M., Lees, A.J., McGeer, P.L. et al. (2004) Genome-wide analysis of the Parkinsonism-dementia complex of Guam. Arch. Neurol., 61, 1889-1897.
46. Poorkaj, P., Tsuang, D., Wijsman, E., Steinbart, E., Garruto, R.M., Craig, U.K., Chapman, N.H., Anderson, L., Bird, T.D., Plato, C.C. et al. (2001) TAU as a susceptibility gene for amyotropic lateral sclerosis-parkinsonism dementia complex of Guam. Arch. Neurol., 58, 1871-1878.
47. Sundar, P.D., Yu, C.E., Sieh, W., Steinbart, E., Garruto, R.M., Oyanagi, K., Craig, U.K., Bird, T.D., Wijsman, E.M., Galasko, D.R. et al. (2007) Two sites in the MAPT region confer genetic risk for Guam ALS/PDC and dementia. Hum. Mol. Genet., 16, 295-306.
48. Galasko, D., Salmon, D., Gamst, A., Olichney, J., Thal, L.J., Silbert, L., Kaye, J., Brooks, P., Adonay, R., Craig, U.K. et al. (2007) Prevalence of dementia in Chamorros on Guam-relationship to age, gender, education, and APOE. Neurology, 68, 1772-1781.
49. Choi, Y., Wijsman, E.M. and Weir, B.S. (2009) Case-control association testing in the presence of unknown relationships. Genet. Epidemiol., in press.
50. Goddard, K.A.B., Yu, C.E., Oshima, J., Miki, T., Nakura, J., Piussan, C., Martin, G.M., Schellenberg, G.D. and Wijsman, E.M. (1996) Toward localization of the Werner Syndrome gene by linkage disequilibrium and ancestral haplotyping: lessons learned from analysis of 35 chromosome 8p11:1-21.1 markers. Am. J. Hum. Genet., 58, 1286-1302.
51. Korbel, J.O., Urban, A.E., Affourtit, J.P., Godwin, B., Grubert, F., Simons, J.F., Kim, P.M., Palejev, D., Carriero, N.J., Du, L. et al. (2007) Paired-end mapping reveals extensive structural variation in the human genome. Science, 318, 420-426.
52. Paisan-Ruiz, C., Jain, S., Evans, E.W., Gilks, W.P., Simon, J., van der Brug, M., de Munain, A.L., Aparicio, S., Gil, A.M., Khan, N. et al. (2004) Cloning of the gene containing mutations that cause PARK8-linked Parkinson's disease. Neuron, 44, 595-600.
53. Zimprich, A., Biskup, S., Leitner, P., Lichtner, P., Farrer, M., Lincoln, S., Kachergus, J., Hulihan, M., Uitti, R.J., Calne, D.B. et al. (2004) Mutations in LRRK2 cause autosomal-dominant Parkinsonism with pleomorphic pathology. Neuron, 44, 601-607.
54. Farrer, L.A., Cupples, L.A., Haines, J.L., Hyman, B., Kukull, W.A., Mayeux, R., Myers, R.H., Pericak-Vance, M.A., Risch, N. and van Duijn, C.M. (1997) Effects of age, sex, and ethnicity on the association between apolipoprotein E genotype and Alzheimer disease. A meta-analysis. APOE and Alzheimer Disease Meta Analysis Consortium. J. Am. Med. Assoc., 278, 1349-1356.
55. Lesage, S., Durr, A., Tazir, M., Lohmann, E., Leutenegger, A.L., Janin, S., Pollak, P. and Brice, A. (2006) LRRK2 G2019S as a cause of Parkinson's disease in North African Arabs. N. Engl. J. Med., 354, 422-423.
56. Gros-Louis, F., Lariviere, R., Gowing, G., Laurent, S., Camu, W., Bouchard, J.P., Meininger, V., Rouleau, G.A. and Julien, J.P. (2004) A frameshift deletion in peripherin gene associated with amyotrophic lateral sclerosis. J. Biol. Chem., 279, 45951-45956.
57. Leung, C.L., He, C.Z., Kaufmann, P., Chin, S.S., Naini, A., Liem, R.K.H., Mitsumoto, H. and Hays, A.P. (2004) A pathogenic peripherin gene mutation in a patient with amyotrophic lateral sclerosis. Brain Pathol., 14, 290-296.
58. Robertson, J., Doroudchi, M.M., Nguyen, M.D., Durham, H.D., Strong, M.J., Shaw, G., Julien, J.P. and Mushynski, W.E. (2003) A neurotoxic peripherin splice variant in a mouse model of ALS. J. Cell Biol., 160, 939-949.
59. Pericak-Vance, M.A., Bass, M.P., Yamaoka, L.H., Gaskell, P.C., Scott, W.K., Terwedow, H.A., Menold, M.M., Conneally, P.M., Small, G.W., Saunders, A.M. et al. (1998) Complete genomic screen in late-onset familial Alzheimer's disease. Neurobiol. Aging, 19, S39-S42.
60. Ertekin-Taner, N. (2007) Genetics of Alzheimer's disease: a centennial review. Neurol. Clin., 25, 611-667.
61. Clark, L.N., Poorkaj, P., Wszolek, Z., Geschwind, D.H., Nasreddine, Z.S., Miller, B., Li, D., Payami, H., Awert, F., Markopoulou, K. et al. (1998) Pathogenic implications of mutations in the tau gene in pallido-ponto-nigral degeneration and related neurodegenerative disorders linked to chromosome 17. Proc. Natl Acad. Sci. USA, 95, 13103-13107.
62. Baker, M., Mackenzie, I.R., Pickering-Brown, S.M., Gass, J., Rademakers, R., Lindholm, C., Snowden, J., Adamson, J., Sadovnick, A.D., Rollinson, S. et al. (2006) Mutations in progranulin cause tau-negative frontotemporal dementia linked to chromosome 17. Nature, 442, 916-919.
63. Hutton, M., Lendon, C.L., Rizzu, P., Baker, M., Froelich, S., Houlden, H., Pickering-Brown, S., Chakraverty, S., Isaacs, A., Grover, A. et al. (1998) Association of missense and 5'-splice-site mutations in tau with the inherited dementia FTDP-17. Nature, 393, 702-705.
64. Bacanu, S.A., Devlin, B. and Roeder, K. (2002) Association studies for quantitative traits in structured populations. Genet. Epidemiol., 22, 78-93.
65. Roeder, K., Bacanu, S.A., Wasserman, L. and Devlin, B. (2006) Using linkage genome scans to improve power of association in genome scans. Am. J. Hum. Genet., 78, 243-252.
66. Hermosura, M.C., Nayakanti, H., Dorovkov, M.V., Calderon, F.R., Ryazanov, A.G., Haymer, D.S. and Garruto, R.M. (2005) A TRPM7 variant shows altered sensitivity to magnesium that may contribute to the pathogenesis of two Guamanian neurodegenerative disorders. Proc. Natl Acad. Sci. USA, 102, 11510-11515.
67. Lynch, D., Wanglund, C., Spathis, R., Chan, C.W., Reiff, D.M., Lum, J.K. and Garruto, R.M. (2008) The contribution of mitochondrial dysfunction to a gene-environment model of Guamanian ALS and PD. Mitochondrion, 8, 109-116.
68. Falchi, M., Forabosco, P., Mocci, E., Borlino, C.C., Picciau, A., Virdis, E., Persico, I., Parracciani, D., Angius, A. and Pirastu, M. (2004) A genomewide search using an original pairwise sampling approach for large genealogies identifies a new locus for total and low-density lipoprotein cholesterol in two genetically differentiated isolates of Sardinia. Am. J. Hum. Genet., 75, 1015-1031.
69. Axenovich, T.I., Zorkoltseva, I.V., Liu, F., Kirichenko, A.V. and Aulchenko, Y.S. (2008) Breaking loops in large complex pedigrees. Hum. Hered., 65, 57-65.
70. Liu, F., Kirichenko, A., Axenovich, T.I., van Duijn, C.M. and Aulchenko, Y.S. (2008) An approach for cutting large and complex pedigrees for linkage analysis. Eur. J. Hum. Genet., 16, 854-860.
71. Margaritte, P., Bonaiti-Pellie, C., King, M.C. and Clerget-Darpoux, F. (1992) Linkage of familial breast cancer to chromosome 17q21 may not be restricted to early-onset disease. Am. J. Hum. Genet., 50, 1231-1234.
72. Broman, K.W., Murray, J.C., Sheffield, V.C., White, R.L. and Weber, J.L. (1998) Comprehensive human genetic maps: individual and sex-specific variation in recombination. Am. J. Hum. Genet., 63, 861-869.
73. Cottingham, R.W., Idury, R.M. and Schaffer, A.A. (1993) Faster sequential genetic linkage computations. Am. J. Hum. Genet., 53, 252-263.
74. Clerget-Darpoux, F., Bonaiti-Pellie, C. and Hochez, J. (1986) Effects of misspecifying genetic parameters in LOD score analysis. Biometrics, 42, 393-399.
75. Ott, J. (1977) Linkage analysis with misclassification at one locus. Clin. Genet., 12, 119-124.
76. Greenberg, D.A. and Hodge, S.E. (1989) Linkage analysis under 'random' and 'genetic' reduced penetrance. Genet. Epidemiol., 6, 259-264.
77. Risch, N. and Giuffra, L. (1992) Model misspecification and multipoint linkage analysis. Hum. Hered., 42, 77-92.
78. Daw, E.W., Thompson, E.A. and Wijsman, E.M. (2000) Bias in multipoint linkage analysis arising from map misspecification. Genet. Epidemiol., 19, 336-380.
79. Fingerlin, T.E., Abecasis, G.R. and Boehnke, M. (2006) Using sex-averaged genetic maps in multipoint linkage analysis when identity-by-descent status is incompletely known. Genet. Epidemiol., 30, 384-396.
80. Halpern, J. and Whittemore, A.S. (1999) Multipoint linkage analysis. A cautionary note. Hum. Hered., 49, 194-196.
81. George, A.W., Wijsman, E.M. and Thompson, E.A. (2005) MCMC multilocus LOD scores: application of a new approach. Hum. Hered., 59, 98-108.
82. Wijsman, E.M., Rothstein, J.H. and Thompson, E.A. (2006) Multipoint linkage analysis with many multiallelic or dense diallelic markers: MCMC provides practical approaches for genome scans on general pedigrees. Am. J. Hum. Genet., 79, 846-858.
83. Lange, K. and Sobel, E. (1991) A random walk method for computing genetic location scores. Am. J. Hum. Genet., 49, 1320-1334.
84. Sobel, E. and Lange, K. (1996) Descent graphs in pedigree analysis: applications to haplotyping, location scores, and marker-sharing statistics. Am. J. Hum. Genet., 58, 1323-1337.
85. Garruto, R.M., Gajdusek, D.C. and Chen, K.M. (1980) Amyotrophic lateral sclerosis among Chamorro migrants from Guam. Ann. Neurol., 8, 612-619.
86. Chapman, N.H. and Wijsman, E.W. (1998) Genome screens using linkage disequilibrium tests: optimal marker characteristics and feasibility. Am. J. Hum. Genet., 63, 1872-1885.
87. Epstein, M.P., Duren, W.L. and Boehnke, M. (2000) Improved inference of relationship for pairs of individuals. Am. J. Hum. Genet., 67, 1219-1231.
88. Milligan, B.G. (2003) Maximum-likelihood estimation of relatedness. Genetics, 163, 1153-1167.
89. Thompson, E.A. (1975) Estimation of pairwise relationships. Ann. Hum. Genet., 39, 173-188.
90. Cotterman, C.W. (1940) A calculus for statistico-genetics. PhD thesis. Ohio State University, Columbia.
91. Bourgain, C., Hoffjan, S., Nicolae, R., Newman, D., Steiner, L., Walker, K., Reynolds, R., Ober, C. and McPeek, M.S. (2003) Novel case-control test in a founder population identifies P-selectin as an atopy-susceptibility locus. Am. J. Hum. Genet., 73, 612-626.
92. Storey, J.D. and Tibshirani, R. (2003) Statistical significance for genomewide studies. Proc. Natl Acad. Sci. USA, 100, 9440-9445.
93. Guo, S.W. and Thompson, E.A. (1989) Analysis of sparse contingency tables: Monte Carlo estimation of exact P-values. Technical Report. Department of Statistics, University of Washington, Seattle.
94. Falush, D., Stephens, M. and Pritchard, J.K. (2003) Inference of population structure using multilocus genotype data: linked loci and correlated allele frequencies. Genetics, 164, 1567-1587.
95. Pritchard, J.K., Stephens, M. and Donnelly, P. (2000) Inference of population structure using multilocus genotype data. Genetics, 155, 945-959.