Meta-analysis of loci associated with age at natural menopause in African-American women

Human Molecular Genetics

Volumn 23, Issue 12, 2014, Pages 3327-3342

  Chen, Christina T L ^a   Liu, Ching Ti ^b,c   Chen, Gary K ^d   Andrews, Jeanette S ^e   Arnold, Alice M ^f   Dreyfus, Jill ^g   Franceschini, Nora ^h   Garcia, Melissa E ^j   Kerr, Kathleen F ^f   Li, Guo ^f   Lohman, Kurt K ^e   Musani, Solomon K ^k   Nalls, Michael A ^l   Raffel, Leslie J ^m   Smith, Jennifer ⁿ   Ambrosone, Christine B ^o   Bandera, Elisa V ^p   Bernstein, Leslie ^q   Britton, Angela ^l   Brzyski, Robert G ^r   Cappola, Anne ^s   Carlson, Christopher S ^a   Couper, David ^h   Deming, Sandra L ^t   Goodarzi, Mark O ^m   Heiss, Gerardo ^h   John, Esther M ^u   Lu, Xiaoning ^b,v   Marchand, Loic Le ^w   Marciante, Kristin ^f   Mcknight, Barbara ^f   Millikan, Robert ^h,i   Nock, Nora L ^x   Olshan, Andrew F ^h,i   Press, Michael F ^d   Vaiyda, Dhananjay ^y   Woods, Nancy F ^f   Taylor, Herman A ^k   Zhao, Wei ⁿ   Zheng, Wei ^t   Evans, Michele K ^l   Harris, Tamara B ^j   Henderson, Brian E ^d   Kardia, Sharon L R ⁿ   Kooperberg, Charles ^a   Liu, Yongmei ^e   Mosley, Thomas H ^k   Psaty, Bruce ^z   Wellons, Melissa ^t   Windham, Beverly G ^k   Zonderman, Alan B ^l   Cupples, L Adrienne ^b,c   Demerath, Ellen W ^g   Haiman, Christopher ^d   Murabito, Joanne M ^c,aa   Rajkovic, Aleksandar ^ab   more..

^a Fred Hutchinson Cancer Research Center   (United States)

^b BOSTON UNIVERSITY   (United States)

^c FRAMINGHAM HEART STUDY   (United States)

^d UNIVERSITY OF SOUTHERN CALIFORNIA   (United States)

^e WAKE FOREST SCHOOL OF MEDICINE   (United States)

^f UNIVERSITY OF WASHINGTON   (United States)

^g UNIVERSITY OF MINNESOTA   (United States)

^h IL   (United States)

ⁱ UNIVERSITY OF NORTH CAROLINA   (United States)

^j Laboratory of Membrane Biochemistry and Biophysics   (United States)

^k UNIVERSITY OF MISSISSIPPI MEDICAL CENTER   (United States)

^l NATIONAL INSTITUTES OF HEALTH   (United States)

^m CEDARS SINAI MEDICAL CENTER   (United States)

ⁿ UNIVERSITY OF MICHIGAN   (United States)

^o ROSWELL PARK CANCER INSTITUTE   (United States)

^p RUTGERS CANCER INSTITUTE OF NEW JERSEY   (United States)

^q CITY OF HOPE NATIONAL MEDICAL CENTER   (United States)

^r Mays Cancer Center at UT Health San Antonio   (United States)

^s UNIVERSITY OF PENNSYLVANIA   (United States)

^t VANDERBILT UNIVERSITY SCHOOL OF MEDICINE   (United States)

^u STANFORD UNIVERSITY SCHOOL OF MEDICINE   (United States)

^v BETH ISRAEL DEACONESS MEDICAL CENTER   (United States)

^w UNIVERSITY OF HAWAII CANCER CENTER   (United States)

^x CASE WESTERN RESERVE UNIVERSITY SCHOOL OF MEDICINE   (United States)

^y JOHNS HOPKINS UNIVERSITY SCHOOL OF MEDICINE   (United States)

^z GROUP HEALTH RESEARCH INSTITUTE   (United States)

^aa BOSTON UNIVERSITY SCHOOL OF MEDICINE   (United States)

^ab UNIVERSITY OF PITTSBURGH SCHOOL OF MEDICINE   (United States)

more..

Author keywords

[No Author keywords available]

Indexed keywords

AFRICAN AMERICAN; ARTICLE; CHROMOSOME 1; CHROMOSOME 19; CHROMOSOME 5; GENE LOCUS; GENE REPLICATION; GENETIC ASSOCIATION; GENETIC VARIABILITY; HUMAN; MENOPAUSE; META ANALYSIS; PRIORITY JOURNAL; UNITED STATES; WOMEN'S HEALTH; AGE; CAUCASIAN; ETHNOLOGY; FEMALE; GENETICS; HUMAN CHROMOSOME;

AFRICAN AMERICANS; AGE FACTORS; CHROMOSOMES, HUMAN; EUROPEAN CONTINENTAL ANCESTRY GROUP; FEMALE; GENETIC LOCI; GENETIC VARIATION; GENOME-WIDE ASSOCIATION STUDY; HUMANS; MENOPAUSE; UNITED STATES;

EID: 84901350518     PISSN: 09646906     EISSN: 14602083     Source Type: Journal    
DOI: 10.1093/hmg/ddu041     Document Type: Article

References (76)

1. Torgerson, D.J., Thomas,R.E. and Reid,D.M.(1997) Mothersand daughters menopausal ages: is there a link? Eur. J. Obstet. Gynecol. Reprod. Biol., 74, 63-66.
2. Parazzini, F. (2007) Determinants of age at menopause in women attending menopause clinics in Italy. Maturitas, 56, 280-287.
3. McKinlay, S.M. (1996) The normal menopause transition: an overview. Maturitas, 23, 137-145.
4. Cooper, G.S. and Sandler, D.P. (1998) Age at natural menopause and mortality. Ann. Epidemiol., 8, 229-235.
5. Jacobsen, B.K., Nilssen, S., Heuch, I. and Kvale, G. (1997) Does age at natural menopause affect mortality from ischemic heart disease? J. Clin. Epidemiol., 50, 475-479.
6. Lindquist, O., Bengtsson, C., Hansson, T. and Roos, B. (1979) Age at menopause and its relation to osteoporosis. Maturitas, 1, 175-181.
7. van Der Voort, D.J., van Der Weijer, P.H. and Barentsen, R. (2003) Early menopause: increased fracture risk at older age. Osteoporos. Int., 14, 525-530.
8. He, L., Tang, X., Li, N., Wu, Y.Q., Wang, J.W., Li, J.R., Zhang, Z.X., Dou, H.D., Liu, J.J., Yu, L.P. et al. (2012) Menopause with cardiovascular disease and its risk factors among rural Chinese women in Beijing: a population-based study. Maturitas, 72, 132-138.
9. van der Schouw, Y.T., van der Graaf, Y., Steyerberg, E.W., Eijkemans, J.C. and Banga, J.D. (1996) Age at menopause as a risk factor for cardiovascular mortality. Lancet, 347, 714-718.
10. Gosden, R.G., Treloar, S.A., Martin, N.G., Cherkas, L.F., Spector, T.D., Faddy, M.J. and Silber, S.J. (2007) Prevalence of premature ovarian failure in monozygotic and dizygotic twins. Hum. Reprod., 22, 610-615.
11. Coulam, C.B. (1982) Premature gonadal failure. Fertil. Steril., 38, 645-655.
12. Murabito, J.M., Yang, Q., Fox, C., Wilson, P.W. and Cupples, L.A. (2005) Heritability of age at natural menopause in the Framingham Heart Study. J. Clin. Endocrinol. Metab., 90, 3427-3430.
13. Morris, D.H., Jones, M.E., Schoemaker, M.J., Ashworth, A. and Swerdlow, A.J. (2011) Familial concordance for age at natural menopause: results from the Breakthrough Generations Study. Menopause, 18, 956-961.
14. de Bruin, J.P., Bovenhuis, H., van Noord, P.A., Pearson, P.L., van Arendonk, J.A., te Velde, E.R., Kuurman, W.W. and Dorland, M. (2001) The role of genetic factors in age at natural menopause. Hum. Reprod., 16, 2014-2018.
15. Mikkelsen, T.F., Graff-Iversen, S., Sundby, J. and Bjertness, E. (2007) Early menopause, association with tobacco smoking, coffee consumption and other lifestyle factors: a cross-sectional study. BMC Public Health, 7, 149.
16. Torgerson, D.J., Avenell, A., Russell, I.T. and Reid, D.M. (1994) Factors associated with onset of menopause in women aged 45-49. Maturitas, 19, 83-92.
17. He, C., Kraft, P., Chen, C., Buring, J.E., Pare, G., Hankinson, S.E., Chanock, S.J., Ridker, P.M., Hunter, D.J. and Chasman, D.I. (2009) Genome-wide association studies identify loci associated with age at menarche and age at natural menopause. Nat. Genet., 41, 724-728.
18. Stolk, L., Zhai, G., van Meurs, J.B., Verbiest, M.M., Visser, J.A., Estrada, K., Rivadeneira, F., Williams, F.M., Cherkas, L., Deloukas, P. et al. (2009) Loci at chromosomes 13, 19 and 20 influence age at natural menopause. Nat. Genet., 41, 645-647.
19. Stolk, L., Perry, J.R.B., Chasman, D.I., He, C., Mangino, M., Sulem, P., Barbalic, M., Broer, L., Byrne, E.M., Ernst, F. et al. (2012) Meta-analyses identify 13 loci associated with age at menopause and highlight DNA repair and immune pathways. Nat. Genet., 44, 260-268.
20. Murray, A., Bennett, C.E., Perry, J.R., Weedon, M.N., Jacobs, P.A., Morris, D.H., Orr, N., Schoemaker, M.J., Jones, M., Ashworth, A. et al. (2010) Common genetic variants are significant risk factors for early menopause: results from the Breakthrough Generations Study. Hum. Mol. Genet., 20, 186-192.
21. Perry, J.R., Corre, T., Esko, T., Chasman, D.I., Fischer, K., Franceschini, N., He, C., Kutalik, Z., Mangino, M., Rose, L.M. et al. (2013) A genome-wide association study of early menopause and the combined impact of identified variants. Hum. Mol. Genet., 22, 1465-1472.
22. Aittomaki, K., Lucena, J.L., Pakarinen, P., Sistonen, P., Tapanainen, J., Gromoll, J., Kaskikari, R., Sankila, E.M., Lehvaslaiho, H., Engel, A.R. et al. (1995) Mutation in the follicle-stimulating hormone receptor gene causes hereditary hypergonadotropic ovarian failure. Cell, 82, 959-968.
23. Laissue, P., Christin-Maitre, S., Touraine, P., Kuttenn, F., Ritvos, O., Aittomaki, K., Bourcigaux, N., Jacquesson, L., Bouchard, P., Frydman, R. et al. (2006) Mutations and sequence variants in GDF9 and BMP15 in patients with premature ovarian failure. Eur. J. Endocrinol., 154, 739-744.
24. Crisponi, L., Deiana, M., Loi, A., Chiappe, F., Uda, M., Amati, P., Bisceglia, L., Zelante, L., Nagaraja, R., Porcu, S. et al. (2001) The putative forkhead transcription factor FOXL2 is mutated in blepharophimosis/ptosis/ epicanthus inversus syndrome. Nat. Genet., 27, 159-166.
25. Rizzolio, F., Bione, S., Sala, C., Goegan, M., Gentile, M., Gregato, G., Rossi, E., Pramparo, T., Zuffardi, O. and Toniolo, D. (2006) Chromosomal rearrangements in Xq and premature ovarian failure: mapping of 25 new cases and review of the literature. Hum. Reprod., 21, 1477-1483.
26. Qin, Y., Choi, Y., Zhao, H., Simpson, J.L., Chen, Z.J. and Rajkovic, A. (2007) NOBOX homeobox mutation causes premature ovarian failure. Am. J. Hum. Genet., 81, 576-581.
27. Hefler, L.A., Grimm, C., Heinze, G., Schneeberger, C., Mueller, M.W., Muendlein, A., Huber, J.C., Leodolter, S. and Tempfer, C.B. (2005) Estrogen-metabolizing gene polymorphisms and age at natural menopause in Caucasian women. Hum. Reprod., 20, 1422-1427.
28. Kevenaar, M.E., Themmen, A.P., Rivadeneira, F., Uitterlinden, A.G., Laven, J.S., van Schoor, N.M., Lips, P., Pols, H.A. and Visser, J.A. (2007)A polymorphism in the AMH type II receptor gene is associated with age at menopause in interaction with parity. Hum. Reprod., 22, 2382-2388.
29. Long, J.R., Shu, X.O., Cai, Q., Cai, H., Gao, Y.T., Jin, F. and Zheng, W. (2006) Polymorphisms of the CYP1B1 gene may be associated with the onset of natural menopause in Chinese women. Maturitas, 55, 238-246.
30. Mitchell, E.S., Farin, F.M., Stapleton, P.L., Tsai, J.M., Tao, E.Y., Smith-DiJulio, K. and Woods, N.F. (2008) Association of estrogen-related polymorphisms with age at menarche, age at final menstrual period, and stages of the menopausal transition. Menopause, 15, 105-111.
31. Tempfer, C.B., Riener, E.K., Keck, C., Grimm, C., Heinze, G., Huber, J.C., Gitsch, G. and Hefler, L.A. (2005) Polymorphisms associated with thrombophilia and vascular homeostasis and the timing of menarche and menopause in 728 white women. Menopause, 12, 325-330.
32. Weel, A.E., Uitterlinden, A.G., Westendorp, I.C., Burger, H., Schuit, S.C., Hofman, A., Helmerhorst, T.J., van Leeuwen, J.P. and Pols, H.A. (1999) Estrogen receptor polymorphism predicts the onset of natural and surgical menopause. J. Clin. Endocrinol. Metab., 84, 3146-3150.
33. He, C., Kraft, P., Chasman, D.I., Buring, J.E., Chen, C., Hankinson, S.E., Pare, G., Chanock, S., Ridker, P.M. and Hunter, D.J. (2010) A large-scale candidate gene association study of age at menarche and age at natural menopause. Hum. Genet., 128, 515-527.
34. Bromberger, J.T., Matthews, K.A., Kuller, L.H., Wing, R.R., Meilahn, E.N. and Plantinga, P. (1997) Prospective study of the determinants of age at menopause. Am. J. Epidemiol., 145, 124-133.
35. Frere, G. (1971) Mean age at menopause and menarche in South Africa. S. Afr. J. Med. Sci., 36, 21-24.
36. Kwawukume, E.Y., Ghosh, T.S. and Wilson, J.B. (1993) Menopausal age of Ghanaian women. Int. J. Gynaecol. Obstet., 40, 151-155.
37. MacMahon, B. and Worcester, J. (1966) Age at menopause. United States-1960-1962. Vital Health Stat. 11, 1-20.
38. Pascall, J.C. and Brown, K.D. (2004) Intramembrane cleavage of ephrinB3 by the human rhomboid family protease, RHBDL2. Biochem. Biophys. Res. Commun., 317, 244-252.
39. Rosenbloom, K.R., Sloan, C.A., Malladi, V.S., Dreszer, T.R., Learned, K., Kirkup, V.M., Wong, M.C., Maddren, M., Fang, R., Heitner, S.G. et al. (2013) ENCODE Data in the UCSC Genome Browser: year 5 update. Nucleic Acids Res., 41, D56-D63.
40. Kilkenny, M.L., De Piccoli, G., Perera, R.L., Labib, K. and Pellegrini, L. (2012) A conserved motif in the C-terminal tail of DNA polymerase alpha tethers primase to the eukaryotic replisome. J. Biol. Chem., 287, 23740-23747.
41. Spencer, K.L., Malinowski, J., Carty, C.L., Franceschini, N., Fernandez-Rhodes, L., Young, A., Cheng, I., Ritchie, M.D., Haiman, C.A., Wilkens, L. et al. (2013) Genetic variation and reproductive timing: African American Women from the Population Architecture Using Genomics and Epidemiology (PAGE) Study. PLoS ONE, 8, e55258.
42. Chen, C.T.L., Fernández-Rhodes, L., Brzyski, R.G., Carlson, C.S., Chen, Z., Heiss, G., North, K.E., Woods, N.F., Rajkovic, A., Kooperberg, C. et al. (2011) Replication of loci influencing ages at menarche and menopause in hispanic women: the Women's health initiative SHARe study. Hum. Mol. Genet., 21, 1419-1432.
43. Schuh-Huerta, S.M., Johnson, N.A., Rosen, M.P., Sternfeld, B., Cedars, M.I. and Reijo Pera, R.A. (2012) Genetic markers of ovarian follicle number and menopause inwomenof multiple ethnicities.Hum. Genet., 131, 1709-1724.
44. Kamiya, T., Kai, W., Tasumi, S., Oka, A., Matsunaga, T., Mizuno, N., Fujita, M., Suetake, H., Suzuki, S., Hosoya, S. et al. (2012)Atrans-species missense SNP in Amhr2 is associated with sex determination in the tiger pufferfish, Takifugu rubripes (fugu). PLoS Genet., 8, e1002798.
45. Knight, P.G. and Glister, C. (2006) TGF-beta superfamily members and ovarian follicle development. Reproduction, 132, 191-206.
46. Freeman, E.W., Sammel, M.D., Lin, H. and Gracia, C.R. (2012) Anti-mullerian hormone as a predictor of time to menopause in late reproductive age women. J. Clin. Endocrinol. Metab., 97, 1673-1680.
47. Prentice, R., Anderson, G., Cummings, S., Freedman, L., Furberg, C., Henderson, M., Johnson, S., Kuller, L., Manson, J., Oberman,A. et al. (1998) Design of the Women's Health Initiative clinical trial and observational study. The Women's Health Initiative Study Group. Control. Clin. Trials, 19, 61-109.
48. Hays, J., Hunt, J.R., Hubbell, F.A., Anderson, G.L., Limacher, M., Allen, C. and Rossouw, J.E. (2003) The Women's Health Initiative recruitment methods and results. Ann. Epidemiol., 13, S18-S77.
49. Li, Y., Willer, C., Sanna, S. and Abecasis, G. (2009) Genotype imputation. Annu. Rev. Genomics Hum. Genet., 10, 387-406.
50. Price, A.L., Patterson, N.J., Plenge, R.M., Weinblatt, M.E., Shadick, N.A. and Reich, D. (2006) Principal components analysis corrects for stratification in genome-wide association studies. Nat. Genet., 38, 904-909.
51. Tang, H., Peng, J., Wang, P. and Risch, N.J. (2005) Estimation of individual admixture: analytical and study design considerations. Genet. Epidemiol., 28, 289-301.
52. FBPP-Investigators. (2002) Multi-center genetic study of hypertension: the Family Blood Pressure Program (FBPP). Hypertension, 39, 3-9.
53. Daniels, P.R., Kardia, S.L., Hanis, C.L., Brown, C.A., Hutchinson, R., Boerwinkle, E. and Turner, S.T. (2004) Familial aggregation of hypertension treatment and control in the Genetic Epidemiology Network of Arteriopathy (GENOA) study. Am. J. Med., 116, 676-681.
54. Pritchard, J.K., Stephens, M. and Donnelly, P. (2000) Inference of population structure using multilocus genotype data. Genetics, 155, 945-959.
55. Purcell, S., Neale, B., Todd-Brown, K., Thomas, L., Ferreira, M.A., Bender, D., Maller, J., Sklar, P., de Bakker, P.I., Daly, M.J. et al. (2007)PLINK:a tool set for whole-genome association and population-based linkage analyses. Am. J. Hum. Genet., 81, 559-575.
56. 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.
57. 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.
58. Falush, D., Stephens, M. and Pritchard, J.K. (2007) Inference of population structure using multilocus genotype data: dominant markers and null alleles. Mol. Ecol. Notes, 7, 574-578.
59. Lettre, G., Palmer, C.D., Young, T., Ejebe, K.G., Allayee, H., Benjamin, E.J., Bennett, F., Bowden, D.W., Chakravarti, A., Dreisbach, A. et al. (2011) Genome-wide association study of coronary heart disease and its risk factors in 8,090 African Americans: the NHLBI CARe Project. PLoS Genet., 7, e1001300.
60. Marchini, J., Howie, B., Myers, S., McVean, G. and Donnelly, P. (2007) A new multipoint method for genome-wide association studies by imputation of genotypes. Nat. Genet., 39, 906-913.
61. Howie, B.N., Donnelly, P. and Marchini, J. (2009) A flexible and accurate genotype imputation method for the next generation of genome-wide association studies. PLoS Genet., 5, e1000529.
62. Musunuru, K., Lettre, G., Young, T., Farlow, D.N., Pirruccello, J.P., Ejebe, K.G., Keating, B.J., Yang, Q., Chen, M.H., Lapchyk, N. et al. (2010) Candidate gene association resource (CARe): design, methods, and proof of concept. Circ. Cardiovasc. Genet., 3, 267-275.
63. Fried, L.P., Borhani, N.O., Enright, P., Furberg, C.D., Gardin, J.M., Kronmal, R.A., Kuller, L.H., Manolio, T.A., Mittelmark, M.B., Newman, A. et al. (1991) The Cardiovascular Health Study: design and rationale. Ann. Epidemiol., 1, 263-276.
64. Tell, G.S., Fried, L.P., Hermanson, B., Manolio, T.A., Newman, A.B. and Borhani, N.O. (1993) Recruitment of adults 65 years and older as participants in the Cardiovascular Health Study. Ann. Epidemiol., 3, 358-366.
65. Browning, B.L. and Browning, S.R. (2009) A unified approach to genotype imputation and haplotype-phase inference for large data sets of trios and unrelated individuals. Am. J. Hum. Genet., 84, 210-223.
66. Psaty, B.M., Lee, M., Savage, P.J., Rutan, G.H., German, P.S. and Lyles, M. (1992) Assessing the use of medications in the elderly: methods and initial experience in the Cardiovascular Health Study. The Cardiovascular Health Study Collaborative Research Group. J. Clin. Epidemiol., 45, 683-692.
67. ARIC-Investigators. (1989) The Atherosclerosis Risk in Communities (ARIC) Study: design and objectives. Am. J. Epidemiol., 129, 687-702.
68. Taylor, H.A. Jr, Wilson, J.G., Jones, D.W., Sarpong, D.F., Srinivasan, A., Garrison, R.J., Nelson, C. and Wyatt, S.B. (2005) Toward resolution of cardiovascular health disparities in African Americans: design and methods of the Jackson Heart Study. Ethnic. Dis., 15, S6-4-S6-17.
69. Bild, D.E., Bluemke, D.A., Burke, G.L., Detrano, R., Diez Roux, A.V., Folsom, A.R., Greenland, P., Jacob, D.R. Jr, Kronmal, R., Liu, K. et al. (2002) Multi-ethnic study of atherosclerosis: objectives and design. Am. J. Epidemiol., 156, 871-881.
70. Newman, B., Moorman, P.G., Millikan, R., Qaqish, B.F., Geradts, J., Aldrich, T.E. and Liu, E.T. (1995) The Carolina Breast Cancer Study: integrating population-based epidemiology and molecular biology. Breast Cancer Res. Treat, 35, 51-60.
71. Kolonel, L.N., Henderson, B.E., Hankin, J.H., Nomura, A.M., Wilkens, L.R., Pike, M.C., Stram, D.O., Monroe, K.R., Earle, M.E. and Nagamine, F.S. (2000) A multiethnic cohort in Hawaii and Los Angeles: baseline characteristics. Am. J. Epidemiol., 151, 346-357.
72. Willer, C.J., Li, Y. and Abecasis, G.R. (2010) METAL: fast and efficient meta-analysis of genomewide association scans. Bioinformatics, 26, 2190-2191.
73. Pruim, R.J., Welch, R.P., Sanna, S., Teslovich, T.M., Chines, P.S., Gliedt, T.P., Boehnke, M., Abecasis, G.R. and Willer, C.J. (2010) LocusZoom: regional visualization of genome-wide association scan results. Bioinformatics, 26, 2336-2337.
74. Encode-Project-Consortium. (2004) The ENCODE (ENCyclopedia Of DNA Elements) Project. Science, 306, 636-640.
75. Geuderman, W.J. and Morrison, J.M. (2006) QUANTO 1.1: a computer program for power and sample size calculations for geneticepidemiology studies. http://hydra.usc.edu/gxe (May 8, 2007 date last accessed).
76. Ernst, J., Kheradpour, P., Mikkelsen, T.S., Shoresh, N., Ward, L.D., Epstein, C.B., Zhang, X., Wang, L., Issner, R., Coyne, M. et al. (2011) Mapping and analysis of chromatin state dynamics in nine human cell types. Nature, 473, 43-49.