[H1] Introduction - University of Edinburgh Research Explorer
Runs of homozygosity: windows into population history and trait architectureFrancisco C. Ceballos1,2, Peter K Joshi3, David W Clark3, Michèle Ramsay1,4 and James F Wilson2,31. Sydney Brenner Institute for Molecular Bioscience, Faculty of Health Sciences, University of the Witwatersrand, Johannesburg, South Africa2. MRC Human Genetics Unit, Institute of Genetics and Molecular Medicine, University of Edinburgh, Western General Hospital, Edinburgh EH4 2XU, Scotland3. Centre for Global Health Research, Usher Institute of Population Health Sciences and Informatics, University of Edinburgh, Teviot Place, Edinburgh EH8 9AG, Scotland 4. Division of Human Genetics, School of Pathology, Faculty of Health Sciences, University of the Witwatersrand, Johannesburg, South AfricaCorrespondence to J.F.W. E-mail: jim.wilson@ed.ac.ukAbstractLong runs of homozygosity (ROH) arise when identical haplotypes are inherited from each parent and thus a long tract of genotypes is homozygous. Cousin marriage or inbreeding gives rise to such autozygosity; however, genome-wide data reveal that ROH are universally common in human genomes even among outbred individuals. The number and length of ROH reflect individual demographic history while the homozygosity burden can be used to investigate the genetic architecture of complex disease. We discuss how to identify ROH in genome-wide microarray and sequence data, their distribution in human populations and application to the understanding of inbreeding depression and disease risk.[H1] IntroductionMating between cousins, or inbreeding ADDIN EN.CITE <EndNote><Cite><Author>Cavalli-Sforza</Author><Year>1978</Year><RecNum>150</RecNum><DisplayText><style face="superscript">1</style></DisplayText><record><rec-number>150</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487159357">150</key></foreign-keys><ref-type name="Book">6</ref-type><contributors><authors><author>Cavalli-Sforza, Luigi L.</author><author>Bodmer, Walter</author></authors></contributors><titles><title>The Genetic of Human Populations</title></titles><dates><year>1978</year></dates><pub-location>San Francisco</pub-location><publisher>W.H.Freeman & Co Ltd</publisher><isbn>0716710188</isbn><urls></urls></record></Cite></EndNote>1, is sometimes considered to be an unusual occurrence in humans; however, such consanguineous marriage is common across the planet. Surveys using genealogical data reveal that at least 10% of the global population (>700 million people) are the offspring of second cousins or closer ADDIN EN.CITE <EndNote><Cite><Author>Bittles</Author><Year>2010</Year><RecNum>22</RecNum><DisplayText><style face="superscript">2</style></DisplayText><record><rec-number>22</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487079810">22</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Bittles, A. H.</author><author>Black, M. L.</author></authors></contributors><titles><title>Consanguinity, human evolution, and complex diseases</title><secondary-title>Proceedings of the National Academy of Sciences of the United States of America</secondary-title><short-title>Consanguinity, human evolution, and complex diseases</short-title></titles><periodical><full-title>Proceedings of the National Academy of Sciences of the United States of America</full-title><abbr-1>Proc. Natl. Acad. Sci. U. S. A.</abbr-1><abbr-2>Proc Natl Acad Sci U S A</abbr-2></periodical><pages>1779-1786</pages><volume>107</volume><number>suppl 1</number><dates><year>2010</year><pub-dates><date>January 26, 2010</date></pub-dates></dates><urls><related-urls><url><style face="underline" font="default" size="100%"> face="underline" font="default" size="100%">. Inbreeding is not distributed evenly around the globe, with higher incidences in areas where consanguinity [G] is favoured culturally such as parts of West and South Asia, but also occurs as a consequence of small population size and endogamy [G] , even if there is random mating. The societal attitudes to cousin marriage are greatly influenced by religious beliefs, with the Quran prohibiting marriages between close relatives, but permission is given for marriage between cousins including double first cousin unions. Even though many examples of consanguinity are cited in biblical texts, the Levitical code also forbids marriage between close kin.Cousins share DNA that they have inherited from their common ancestors and thus the offspring of cousin marriage may inherit identical chromosomal segments from both parents. The availability of denser genome-wide microsatellite scans in the mid-1990s led to the discovery of uninterrupted long runs of homozygous genotypes (known as runs of homozygosity [G] (ROH)), the hallmark of these autozygous [G] segments inherited from a recent common ancestor ADDIN EN.CITE <EndNote><Cite><Author>Broman</Author><Year>1999</Year><RecNum>25</RecNum><DisplayText><style face="superscript">3</style></DisplayText><record><rec-number>25</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487079810">25</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Broman, K. W.</author><author>Weber, J. L.</author></authors></contributors><titles><title>Long homozygous chromosomal segments in reference families from the centre d'Etude du polymorphisme humain</title><secondary-title>American Journal of Human Genetics</secondary-title><short-title>Long homozygous chromosomal segments in reference families from the centre d'Etude du polymorphisme humain</short-title></titles><periodical><full-title>American Journal of Human Genetics</full-title><abbr-1>Am. J. Hum. Genet.</abbr-1><abbr-2>Am J Hum Genet</abbr-2></periodical><pages>1493-500</pages><volume>65</volume><dates><year>1999</year></dates><label>Broman1999</label><urls><related-urls><url><style face="underline" font="default" size="100%"> face="underline" font="default" size="100%">. Members of two families recruited to construct the first human genetic maps — of Venezuelan and Old Order Amish ancestry — carried 4–16 ROH typically ~1.5–30 Mb in length, the most extreme individual having a total of ~195 Mb in ROH, consistent with close inbreeding ADDIN EN.CITE <EndNote><Cite><Author>Broman</Author><Year>1999</Year><RecNum>25</RecNum><DisplayText><style face="superscript">3</style></DisplayText><record><rec-number>25</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487079810">25</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Broman, K. W.</author><author>Weber, J. L.</author></authors></contributors><titles><title>Long homozygous chromosomal segments in reference families from the centre d'Etude du polymorphisme humain</title><secondary-title>American Journal of Human Genetics</secondary-title><short-title>Long homozygous chromosomal segments in reference families from the centre d'Etude du polymorphisme humain</short-title></titles><periodical><full-title>American Journal of Human Genetics</full-title><abbr-1>Am. J. Hum. Genet.</abbr-1><abbr-2>Am J Hum Genet</abbr-2></periodical><pages>1493-500</pages><volume>65</volume><dates><year>1999</year></dates><label>Broman1999</label><urls><related-urls><url><style face="underline" font="default" size="100%"> face="underline" font="default" size="100%">. More unexpected was the fact that, despite the relatively sparse and imperfect maps, 20% of the 100 individuals outside these two families (all Utah Mormons) carried at least one homozygous segment — ROH were thus likely to be common in human populations. It becomes clear why ROH are common when we consider that an individual today is predicted to have half a billion (229) ancestors 29 generations ago (c1100, one generation after William the Conqueror), more than the estimated world population of ~310 million at that time ADDIN EN.CITE <EndNote><Cite><Author>McEvedy</Author><Year>1978</Year><RecNum>195</RecNum><DisplayText><style face="superscript">4,5</style></DisplayText><record><rec-number>195</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1488465157">195</key></foreign-keys><ref-type name="Book">6</ref-type><contributors><authors><author>Colin. McEvedy
Richard M. Jones</author></authors></contributors><titles><title>Atlas of world Population History</title></titles><dates><year>1978</year></dates><pub-location>New York</pub-location><publisher>Facts On File</publisher><urls></urls></record></Cite><Cite><Author>Biraben</Author><Year>1980</Year><RecNum>198</RecNum><record><rec-number>198</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1488466746">198</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Biraben, Jean-Noel</author></authors></contributors><titles><title>An essay concerning mankind's demographic evolution</title><secondary-title>Journal of human Evolution</secondary-title></titles><periodical><full-title>Journal of Human Evolution</full-title><abbr-1>J. Hum. Evol.</abbr-1><abbr-2>J Hum Evol</abbr-2></periodical><pages>655-663</pages><volume>9</volume><dates><year>1980</year></dates><urls></urls></record></Cite></EndNote>4,5. This ancestor paradox is solved by the fact that many of the ancestors are the same people (known as pedigree collapse ADDIN EN.CITE <EndNote><Cite><Author>Gunderson</Author><Year>1980</Year><RecNum>192</RecNum><DisplayText><style face="superscript">6</style></DisplayText><record><rec-number>192</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1488464922">192</key></foreign-keys><ref-type name="Book">6</ref-type><contributors><authors><author>Gunderson, Robert C.</author></authors></contributors><titles><title>Connecting your pedigree into royal, noble and medieval families</title></titles><dates><year>1980</year></dates><pub-location>Salt Lake City</pub-location><publisher>Genealogical Society of Utah</publisher><urls></urls></record></Cite></EndNote>6). In most cases, given broad- and fine-scale human population genetic structure and limited effective population sizes [G] , ancestors will be shared more recently in time than the 12th centuryPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5LZWxsZXI8L0F1dGhvcj48WWVhcj4yMDExPC9ZZWFyPjxS
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ADDIN EN.CITE.DATA 7: we are all inbred to some degree and ROH capture this aspect of our individual demographic histories [G]. To this end they can even be analysed by free online utilities for genetic genealogists who have purchased direct-to-consumer genome scans. We do not inherit DNA from all our pedigree ancestors at this remove of generations ADDIN EN.CITE <EndNote><Cite><Author>Donnelly</Author><Year>1983</Year><RecNum>205</RecNum><DisplayText><style face="superscript">8</style></DisplayText><record><rec-number>205</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1488617248">205</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Donnelly, Kevin P.</author></authors></contributors><titles><title>The probability that related individuals share some section of genome identical by descent</title><secondary-title>Theoretical Population Biology</secondary-title></titles><periodical><full-title>Theoretical Population Biology</full-title></periodical><pages>34-63</pages><volume>23</volume><number>1</number><dates><year>1983</year></dates><isbn>00405809</isbn><urls></urls><electronic-resource-num>10.1016/0040-5809(83)90004-7</electronic-resource-num></record></Cite></EndNote>8; however, we have to inherit DNA from some of them and as the number of genealogical ancestors doubles every generation, eventually everyone has shared genetic ancestors, between 300–1400 BC, depending on assumptions about migration ADDIN EN.CITE <EndNote><Cite><Author>Rohde</Author><Year>2004</Year><RecNum>218</RecNum><DisplayText><style face="superscript">9</style></DisplayText><record><rec-number>218</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1488874092">218</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Rohde, D. L.</author><author>Olson, S.</author><author>Chang, J. T.</author></authors></contributors><auth-address>Department of Brain and Cognitive Sciences, Massachusetts Institute of Technology, Cambridge, Massachusetts 02139, USA.</auth-address><titles><title>Modelling the recent common ancestry of all living humans</title><secondary-title>Nature</secondary-title></titles><periodical><full-title>Nature</full-title></periodical><pages>562-6</pages><volume>431</volume><number>7008</number><keywords><keyword>*Computer Simulation</keyword><keyword>Emigration and Immigration</keyword><keyword>Female</keyword><keyword>Geography</keyword><keyword>Humans</keyword><keyword>Male</keyword><keyword>Monte Carlo Method</keyword><keyword>*Pedigree</keyword><keyword>*Phylogeny</keyword><keyword>Population Density</keyword><keyword>Population Dynamics</keyword><keyword>Reproduction</keyword><keyword>Time Factors</keyword></keywords><dates><year>2004</year><pub-dates><date>Sep 30</date></pub-dates></dates><isbn>1476-4687 (Electronic)
0028-0836 (Linking)</isbn><accession-num>15457259</accession-num><urls><related-urls><url><style face="underline" font="default" size="100%">. It has been known for over a century ADDIN EN.CITE <EndNote><Cite><Author>Garrod</Author><Year>1902</Year><RecNum>206</RecNum><DisplayText><style face="superscript">10</style></DisplayText><record><rec-number>206</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1488617512">206</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Garrod, Archibald E.</author></authors></contributors><titles><title>The incidence of alkaptonuria: a study in chemical individuality</title><secondary-title>Lancet Infectious Diseases</secondary-title></titles><periodical><full-title>Lancet Infectious Diseases</full-title><abbr-1>Lancet Infect. Dis.</abbr-1><abbr-2>Lancet Infect Dis</abbr-2></periodical><pages>1616-1620</pages><volume>2</volume><dates><year>1902</year></dates><urls></urls></record></Cite></EndNote>10 that inbreeding increases the incidence of recessive disease, and the frequency of homozygotes is increased in relation to the inbreeding level in the population. The long ROH in inbred individuals reveal the full, harmful effects of recessive deleterious variants present in the ROH, e.g. to cause Mendelian diseases such as Tay–Sachs. Inbreeding usually leads to decreases in the vigour and reproductive fitness of offspring — known as inbreeding depression [G] — as first noted by Charles Darwin in plants (Box 1) and seen for numerous fitness-related traits in animalsPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5Ib2ZmbWFuPC9BdXRob3I+PFllYXI+MjAxNDwvWWVhcj48
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ADDIN EN.CITE.DATA 11,12.In this Review we focus on the analyses of human ROH (rather than single-marker inbreeding coefficients) and their contributions to the understanding of human demographic history and in deciphering the genetic architecture [G] of complex disease. We do not focus on Mendelian conditions or human knockouts. We discuss methodological considerations regarding the identification of ROH in microarray and sequence datasets, the distribution of ROH of different lengths across the genome and the globe; and their relation to pedigree. We review the burgeoning literature on the influence of ROH on disease risk and quantitative traits, and what has been learned about inbreeding depression in humans. We conclude with some recommendations for the assessment of ROH and highlight future research questions.[H1] Origins of ROH and inbreeding depressionROH arise when two copies of an ancestral haplotype [G] are brought together in an individual: longer haplotypes inherited from recent common ancestors or shorter haplotypes from distant ones (background relatedness). Short ROH characterized by strong linkage disequilibrium (LD) among markers are not always considered autozygous, but nevertheless are due to mating of distantly related individuals. Different population histories give rise to divergent distributions of long and short ROH (Fig. 1). The ROH complement of outbred populations is related to their effective population size, with smaller populations tending to have more ROH and larger populations fewer. Admixed [G] populations, on account of their more distant shared ancestry across the two or more ancestral populations, have fewer ROH than their respective parental populations. Consanguineous communities, on the other hand, have much longer ROH than seen in outbred populations due to very recent pedigree inbreeding loops [G] , whereas populations that have undergone a population bottleneck [G] carry a greater number of shorter ROH than cosmopolitan populations [G] , reflecting deeper parental relatedness. Finally, populations with both reduced effective population size in the past and recent inbreeding have the greatest burden of ROH.The causal mechanism for inbreeding depression is only partly understood, but empirical evidence in a number of species suggests that it is mostly due to increased homozygosity for (partially) recessive detrimental mutations maintained at low frequency in populations by mutation–selection balance, although the contribution of some loci with heterozygote advantage (overdominance [G] ) maintained at intermediate frequencies by balancing selection [G] cannot be discarded ADDIN EN.CITE <EndNote><Cite><Author>Charlesworth</Author><Year>2009</Year><RecNum>36</RecNum><DisplayText><style face="superscript">13</style></DisplayText><record><rec-number>36</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487079810">36</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Charlesworth, D.</author><author>Willis, J. H.</author></authors></contributors><titles><title>The genetics of inbreeding depression</title><secondary-title>Nature Reviews Genetics</secondary-title><short-title>FUNDAMENTAL CONCEPTS IN GENETICS The genetics of inbreeding depression</short-title></titles><periodical><full-title>Nature Reviews Genetics</full-title><abbr-1>Nat. Rev. Genet.</abbr-1><abbr-2>Nat Rev Genet</abbr-2></periodical><pages>783-796</pages><volume>10</volume><number>11</number><dates><year>2009</year><pub-dates><date>Nov</date></pub-dates></dates><isbn>1471-0056</isbn><accession-num>WOS:000270915400014</accession-num><urls><related-urls><url><Go to ISI>://WOS:000270915400014</url></related-urls></urls><electronic-resource-num>10.1038/nrg2664</electronic-resource-num></record></Cite></EndNote>13. When dominant alleles at some loci decrease the trait value while others increase it, we do not expect any association with genome-wide homozygosity. However, if on average across all causal loci, dominance [G] is biased in one direction, for instance to decrease the trait, we will see such an association. Such directional dominance [G] arises due to directional selection in evolutionary fitness-related traits.Empirical studies ADDIN EN.CITE <EndNote><Cite><Author>Szpiech</Author><Year>2013</Year><RecNum>209</RecNum><DisplayText><style face="superscript">14</style></DisplayText><record><rec-number>209</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1488617963">209</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Szpiech, Z. A.</author><author>Xu, J.</author><author>Pemberton, T. J.</author><author>Peng, W.</author><author>Zollner, S.</author><author>Rosenberg, N. A.</author><author>Li, J. Z.</author></authors></contributors><auth-address>Department of Bioengineering and Therapeutic Sciences, University of California, San Francisco, San Francisco, CA 94158, USA. zachary.szpiech@ucsf.edu</auth-address><titles><title>Long runs of homozygosity are enriched for deleterious variation</title><secondary-title>American Journal of Human Genetics</secondary-title></titles><periodical><full-title>American Journal of Human Genetics</full-title><abbr-1>Am. J. Hum. Genet.</abbr-1><abbr-2>Am J Hum Genet</abbr-2></periodical><pages>90-102</pages><volume>93</volume><number>1</number><keywords><keyword>Alleles</keyword><keyword>Computational Biology/methods</keyword><keyword>Consanguinity</keyword><keyword>Exome</keyword><keyword>Genetics, Population/*methods</keyword><keyword>*Genome, Human</keyword><keyword>*Genomic Structural Variation</keyword><keyword>Heterozygote</keyword><keyword>*Homozygote</keyword><keyword>Humans</keyword><keyword>Mutation, Missense</keyword><keyword>Polymorphism, Single Nucleotide</keyword><keyword>Predictive Value of Tests</keyword></keywords><dates><year>2013</year><pub-dates><date>Jul 11</date></pub-dates></dates><isbn>1537-6605 (Electronic)
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ADDIN EN.CITE.DATA 15, although this is expected given the typically lower allele frequencies of deleterious variants compared to non-deleterious variants. Inbreeding increases the probability that a variant will be found in a homozygous state, so ROH are enriched for homozygotes at all allele frequencies. This enrichment is particularly strong for rare variants since a variant at frequency p is homozygous at frequency p2 outside ROH, and at frequency p inside ROH (where p is the population frequency of the allele). Homozygotes thus occur (1/p) times more frequently inside ROH, so lower frequency variants (including deleterious variants) are more strongly enriched. Theory also predicts that very strong inbreeding will in fact purge deleterious recessive alleles from the population as more copies are found in a homozygous state, and this has been observed in mountain and eastern lowland gorillasPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5YdWU8L0F1dGhvcj48WWVhcj4yMDE1PC9ZZWFyPjxSZWNO
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ADDIN EN.CITE.DATA 16 but not in human genome dataPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5TY290dDwvQXV0aG9yPjxZZWFyPjIwMTY8L1llYXI+PFJl
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ADDIN EN.CITE.DATA 17. [H1] Methodological considerationsROH calling requires high-density genome-wide scan data, now overwhelmingly from single nucleotide polymorphism (SNP) microarrays, but ROH analysis of short-read sequencing of the entire genome or exome will become more common as the price of these technologies decreases. A number of factors influence the quality of ROH calling, including the marker density, their distribution across the genome, the quality of the genotype calling (including error rates), and minor-allele frequencies. Microarray data are considered gold standard with very low error rates (typically <0.1%); however, the content usually comprises ~1 million SNPs with allele frequencies >5%, chosen to best represent haplotype structure. Whole-genome sequencing (WGS), on the other hand, assays every variant, irrespective of allele frequency, although the low coverage often employed to maximize the number of individual genomes sequenced and hence power for association, means that rare SNPs are called significantly less often, with higher error rates, than common SNPs. Hence parameters of calling algorithms require tuning to the characteristics of the underlying data, and particular care must be taken with centromeres, duplications and other difficult regions. There are two major methods for identifying ROH: observational genotype-counting ADDIN EN.CITE <EndNote><Cite><Author>Purcell</Author><Year>2007</Year><RecNum>116</RecNum><DisplayText><style face="superscript">18</style></DisplayText><record><rec-number>116</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487079810">116</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Purcell, S.</author><author>Neale, B.</author><author>Todd-Brown, K.</author><author>Thomas, L.</author><author>Ferreira, M. A. R.</author><author>Bender, D.</author><author>Maller, J.</author><author>Sklar, P.</author><author>de Bakker, P. I. W.</author><author>Daly, M. J.</author><author>Sham, P. C.</author></authors></contributors><titles><title>PLINK: A tool set for whole-genome association and population-based linkage analyses</title><secondary-title>American Journal of Human Genetics</secondary-title><short-title>PLINK: A tool set for whole-genome association and population-based linkage analyses</short-title></titles><periodical><full-title>American Journal of Human Genetics</full-title><abbr-1>Am. J. Hum. Genet.</abbr-1><abbr-2>Am J Hum Genet</abbr-2></periodical><pages>559-575</pages><volume>81</volume><number>3</number><dates><year>2007</year><pub-dates><date>Sep</date></pub-dates></dates><isbn>0002-9297</isbn><accession-num>WOS:000249128200012</accession-num><urls><related-urls><url><Go to ISI>://WOS:000249128200012</url><url> and model-basedPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5QZW1iZXJ0b248L0F1dGhvcj48WWVhcj4yMDEyPC9ZZWFy
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ADDIN EN.CITE.DATA 19. [H3] Observational approaches. Algorithms, such as that implemented in PLINK ADDIN EN.CITE <EndNote><Cite><Author>Purcell</Author><Year>2007</Year><RecNum>116</RecNum><DisplayText><style face="superscript">18</style></DisplayText><record><rec-number>116</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487079810">116</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Purcell, S.</author><author>Neale, B.</author><author>Todd-Brown, K.</author><author>Thomas, L.</author><author>Ferreira, M. A. R.</author><author>Bender, D.</author><author>Maller, J.</author><author>Sklar, P.</author><author>de Bakker, P. I. W.</author><author>Daly, M. J.</author><author>Sham, P. C.</author></authors></contributors><titles><title>PLINK: A tool set for whole-genome association and population-based linkage analyses</title><secondary-title>American Journal of Human Genetics</secondary-title><short-title>PLINK: A tool set for whole-genome association and population-based linkage analyses</short-title></titles><periodical><full-title>American Journal of Human Genetics</full-title><abbr-1>Am. J. Hum. Genet.</abbr-1><abbr-2>Am J Hum Genet</abbr-2></periodical><pages>559-575</pages><volume>81</volume><number>3</number><dates><year>2007</year><pub-dates><date>Sep</date></pub-dates></dates><isbn>0002-9297</isbn><accession-num>WOS:000249128200012</accession-num><urls><related-urls><url><Go to ISI>://WOS:000249128200012</url><url>, scan each chromosome by moving a window of fixed size along their length in search of stretches of consecutive homozygous SNPs. ROH are called by first calculating the proportion of completely homozygous windows that encompass that SNP. If this proportion is higher than a defined threshold, the SNP is designated as being in an ROH. A variable number of heterozygous or missing SNPs per window can be specified in order to tolerate genotyping errors and failures, as well as rare new mutation events. Finally, an ROH is called if the number of consecutive SNPs in a homozygous segment exceeds a predefined threshold in terms of SNP number and/or covered chromosomal length. The simplicity of the PLINK approach allows distributed applications in large consortiabXBlcmlhbCBDb2xsZWdlIExvbmRvbiwgTm9yZm9sayBQbGFjZSwgTG9uZG9uIFcyIDFQRywgVUsu
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ADDIN EN.CITE.DATA 20 and SNP data may be pruned for LD if desired before ROH calling. Haplotype-matching algorithms (e.g. Germline ADDIN EN.CITE <EndNote><Cite><Author>Gusev</Author><Year>2009</Year><RecNum>59</RecNum><DisplayText><style face="superscript">21</style></DisplayText><record><rec-number>59</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487079810">59</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Gusev, A.</author><author>Lowe, J. K.</author><author>Stoffel, M.</author><author>Daly, M. J.</author><author>Altshuler, D.</author><author>Breslow, J. L.</author><author>Friedman, J. M.</author><author>Pe'er, I.</author></authors></contributors><auth-address>Department of Computer Science, Columbia University, New York, New York 10027, USA. gusev@cs.columbia.edu</auth-address><titles><title>Whole population, genome-wide mapping of hidden relatedness</title><secondary-title>Genomic Research</secondary-title><short-title>Whole population, genome-wide mapping of hidden relatedness</short-title></titles><periodical><full-title>Genomic Research</full-title></periodical><pages>318-26</pages><volume>19</volume><number>2</number><keywords><keyword>Algorithms</keyword><keyword>Chromosome Mapping/ methods</keyword><keyword>Consanguinity</keyword><keyword>Family Relations</keyword><keyword>Genetics, Population/methods</keyword><keyword>Haplotypes</keyword><keyword>Humans</keyword><keyword>Inheritance Patterns/ genetics</keyword><keyword>Models, Genetic</keyword><keyword>Phylogeny</keyword><keyword>Polymorphism, Single Nucleotide</keyword><keyword>Sensitivity and Specificity</keyword></keywords><dates><year>2009</year></dates><pub-location>United States</pub-location><isbn>1088-9051 (Print) 1088-9051 (Linking)</isbn><accession-num>18971310</accession-num><urls><related-urls><url><style face="underline" font="default" size="100%">) for calculation of identity-by-descent [G] (IBD) can also be used to identify ROH, as a special case of IBD within an individual. [H3] Model-based approaches. An alternative, computationally expensive approach implemented in Beagle uses hidden Markov models (HMMs) to account for background levels of LDPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5Ccm93bmluZzwvQXV0aG9yPjxZZWFyPjIwMTA8L1llYXI+
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ADDIN EN.CITE.DATA 22. However, tests on simulated data showed that PLINK outperformed Germline and Beagle in detecting ROHPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5Ib3dyaWdhbjwvQXV0aG9yPjxZZWFyPjIwMTE8L1llYXI+
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ADDIN EN.CITE.DATA 23. Further likelihood-based approaches use the log of the ratio of the probabilities of the genotype data under the hypotheses of autozygosity and non-autozygosity (incorporating population-specific allele frequency estimates) to infer the homozygosity status of sliding windows in each individualPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5QZW1iZXJ0b248L0F1dGhvcj48WWVhcj4yMDEyPC9ZZWFy
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ADDIN EN.CITE.DATA 19. A population-specific threshold was defined from these log-odds scores, above which ROH are called. Gaussian kernel density estimates of the genome-wide log-odds scores revealed two modes in each population and the local minimum was used as the threshold in each case. The distribution of ROH lengths was also modelled as a mixture of three Gaussian distributions, classifying ROH into size classes: very short ROHs (tens to hundreds of kb) reflecting LD patterns; intermediate ROH (hundreds of kb to 2?Mb) that result from background relatedness owing to genetic drift; and long ROH (over 1–2?Mb) arising from recent parental relatednessPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5QZW1iZXJ0b248L0F1dGhvcj48WWVhcj4yMDEyPC9ZZWFy
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ADDIN EN.CITE.DATA 19. Despite providing increased sensitivity in the detection of shorter ROH, the need to estimate allele frequencies is a limitation of this approach — now implemented in the Garlic software ADDIN EN.CITE <EndNote><Cite><Author>Szpiech</Author><Year>2017</Year><RecNum>219</RecNum><DisplayText><style face="superscript">24</style></DisplayText><record><rec-number>219</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1488874193">219</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Szpiech, Z. A.</author><author>Blant, A.</author><author>Pemberton, T. J.</author></authors></contributors><auth-address>Department of Bioengineering and Therapeutic Sciences, Sciences, University of California-San Francisco, San Francisco, CA, USA 94158.
Department of Biochemistry and Medical Genetics, University of Manitoba, Winnipeg, MB, Canada R3E 0J9.</auth-address><titles><title>GARLIC: Genomic Autozygosity Regions Likelihood-based Inference and Classification</title><secondary-title>Bioinformatics</secondary-title></titles><periodical><full-title>Bioinformatics</full-title><abbr-1>Bioinformatics</abbr-1><abbr-2>Bioinformatics</abbr-2></periodical><dates><year>2017</year><pub-dates><date>Feb 16</date></pub-dates></dates><isbn>1367-4811 (Electronic)
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ADDIN EN.CITE.DATA 27; however, analysis requires the very large mapped sequence read (bam) files. Further HMM methods for sequence data include BCFtools/ROH ADDIN EN.CITE <EndNote><Cite><Author>Narasimhan</Author><Year>2016</Year><RecNum>210</RecNum><DisplayText><style face="superscript">28</style></DisplayText><record><rec-number>210</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1488618138">210</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Narasimhan, V.</author><author>Danecek, P.</author><author>Scally, A.</author><author>Xue, Y.</author><author>Tyler-Smith, C.</author><author>Durbin, R.</author></authors></contributors><auth-address>Wellcome Trust Sanger Institute, Hinxton and.
Department of Genetics, University of Cambridge, Cambridge, UK.</auth-address><titles><title>BCFtools/RoH: a hidden Markov model approach for detecting autozygosity from next-generation sequencing data</title><secondary-title>Bioinformatics</secondary-title></titles><periodical><full-title>Bioinformatics</full-title><abbr-1>Bioinformatics</abbr-1><abbr-2>Bioinformatics</abbr-2></periodical><pages>1749-51</pages><volume>32</volume><number>11</number><dates><year>2016</year><pub-dates><date>Jun 01</date></pub-dates></dates><isbn>1367-4811 (Electronic)
1367-4803 (Linking)</isbn><accession-num>26826718</accession-num><urls><related-urls><url>, which has similarly low error rates and can use much smaller variant call format (vcf) files, which contain only the SNP genotypes and quality scores. High-depth WGS holds the promise of the most accurate ROH detection, and will allow assessment of the contribution of very short ROH to inbreeding depression.[H1] Distribution of runs of homozygosity[H3] Runs of homozygosity are ubiquitous. A survey using ~700,000 SNP microarray genotypes for 209 HapMap individuals revealed for the first time how widespread megabase-scale ROH were, even among outbred individuals ADDIN EN.CITE <EndNote><Cite><Author>Gibson</Author><Year>2006</Year><RecNum>55</RecNum><DisplayText><style face="superscript">29</style></DisplayText><record><rec-number>55</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487079810">55</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Gibson, J.</author><author>Morton, N. E.</author><author>Collins, A.</author></authors></contributors><titles><title>Extended tracts of homozygosity in outbred human populations</title><secondary-title>Human Molecular Genetics</secondary-title><short-title>Extended tracts of homozygosity in outbred human populations</short-title></titles><periodical><full-title>Human Molecular Genetics</full-title><abbr-1>Hum. Mol. Genet.</abbr-1><abbr-2>Hum Mol Genet</abbr-2></periodical><pages>789-795</pages><volume>15</volume><dates><year>2006</year></dates><label>Gibson2006</label><urls><related-urls><url><style face="underline" font="default" size="100%"> face="underline" font="default" size="100%">. However, different continental populations have contrasting burdens: Africans generally have fewer ROH, reflecting their larger effective population size. Again, this survey identified cryptically inbred outliers: a Mormon from Utah and two Japanese from Tokyo. Further studies verified these findings in European AmericansPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5TaW1vbi1TYW5jaGV6PC9BdXRob3I+PFllYXI+MjAwNzwv
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ADDIN EN.CITE.DATA 3,34. Analysis of >3 million SNPs in the HapMap populations allowed identification of ROH down to 100?kb in length, which are dramatically more numerous: each individual carries hundreds to thousands of these, which in total comprise 400–500?Mb of the genomes of cosmopolitan Europeans and East Asians, but only 160?Mb in Yoruba from NigeriaPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5JbnRlcm5hdGlvbmFsIEhhcE1hcDwvQXV0aG9yPjxZZWFy
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ADDIN EN.CITE.DATA 33. Thus, such short ROH account for more of the total sum of ROH than ROH >1?Mb, even for inbred individuals.[H3] Correlation with pedigree inbreeding. The degree of individual inbreeding is measured using the inbreeding coefficient [G] (F), the probability that an individual receives two alleles that are identical-by-descent at a given locus ADDIN EN.CITE <EndNote><Cite><Author>Wright</Author><Year>1922</Year><RecNum>211</RecNum><DisplayText><style face="superscript">36</style></DisplayText><record><rec-number>211</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1488618288">211</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Sewall Wright</author></authors></contributors><titles><title>Coefficients of Inbreeding and relationship</title><secondary-title>The American Naturalist</secondary-title></titles><periodical><full-title>The American Naturalist</full-title><abbr-1>Amer. Naturalist</abbr-1></periodical><pages>330-338</pages><volume>56</volume><dates><year>1922</year></dates><urls></urls></record></Cite></EndNote>36, which is also the expected proportion of the genome that is autozygous, e.g. F=0.0625 for the offspring of first cousins. The genomic inbreeding coefficient [G] , FROH, measures the actual proportion of the autosomal genome that is autozygous — defined as the sum total length of ROH (SROH) over a specified minimum length threshold, as a proportion of the total genome lengthPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5NY1F1aWxsYW48L0F1dGhvcj48WWVhcj4yMDA4PC9ZZWFy
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ADDIN EN.CITE.DATA 35. Another useful measure of ROH is the total number of ROH (NROH). FROH captures the total inbreeding coefficient of the individual irrespective of pedigree accuracy or depth (or absence), within the resolution of the dataset available (and hence the size of ROH that can be called). Early studies revealed that offspring of first cousins with autosomal recessive disease had a mean FROH of 11%, substantially higher than predicted, probably due to generations of consanguineous marriagePEVuZE5vdGU+PENpdGU+PEF1dGhvcj5Xb29kczwvQXV0aG9yPjxZZWFyPjIwMDY8L1llYXI+PFJl
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ADDIN EN.CITE PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5Xb29kczwvQXV0aG9yPjxZZWFyPjIwMDY8L1llYXI+PFJl
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ADDIN EN.CITE.DATA 37. Analysis of a broader spectrum of parental relatedness using accurate pedigrees from an isolated population demonstrated that FROH calculated using ROH >1.5?Mb in length correlated most strongly (r=0.86) with inbreeding coefficients from 6-generation pedigrees (Fped)PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5NY1F1aWxsYW48L0F1dGhvcj48WWVhcj4yMDA4PC9ZZWFy
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ADDIN EN.CITE.DATA 35. In fact, across diverse samples, population mean Fped also correlates well with FROH using ROH >5?Mb (r=0.87), but not with FROH calculated from ROH <5?MbPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5LaXJpbjwvQXV0aG9yPjxZZWFyPjIwMTA8L1llYXI+PFJl
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ADDIN EN.CITE.DATA 38.[H3] Global distribution. The distribution of ROH across worldwide populations is structured at many scales from continental to tribalPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5LaXJpbjwvQXV0aG9yPjxZZWFyPjIwMTA8L1llYXI+PFJl
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ADDIN EN.CITE.DATA 41, but also the Selkup of Siberia — have an increased mean SROH and usually increased variance as well. As the relatively small number of very long ROH arising from the recent inbreeding loops influences the sum of ROH much more than the total number, these populations display a ‘right shift’ in the NROH versus SROH graph away from the trend line (Fig. 2b). Long tails in the distributions of SROH, or increased means, are also seen (Fig. 2a). A second class includes numerous native populations from across the world with shared parental ancestry arising from isolation and endogamy over many generations, but comparatively little recent inbreeding. Such individuals carry few long ROH, but have substantial enrichment for ROH in the 2–5?Mb length, leading to a relatively high NROH, and include Papua New Guinean Highlanders, the Koryak and Chukchi of Siberia, the Pulliyar, Kurumba and Piramalai Kallar castes of Southern India, unadmixed Greenlanders and Athapaskans of North America, the click-speaking Hadza hunter-gatherers of Tanzania (Fig 2ab) and the Onge of the Andaman Islands in the Bay of BengalPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5Kb25lczwvQXV0aG9yPjxZZWFyPjIwMTU8L1llYXI+PFJl
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ADDIN EN.CITE.DATA 20,44, e.g. Amish (in whom ROH were first discovered ADDIN EN.CITE <EndNote><Cite><Author>Broman</Author><Year>1999</Year><RecNum>25</RecNum><DisplayText><style face="superscript">3</style></DisplayText><record><rec-number>25</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487079810">25</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Broman, K. W.</author><author>Weber, J. L.</author></authors></contributors><titles><title>Long homozygous chromosomal segments in reference families from the centre d'Etude du polymorphisme humain</title><secondary-title>American Journal of Human Genetics</secondary-title><short-title>Long homozygous chromosomal segments in reference families from the centre d'Etude du polymorphisme humain</short-title></titles><periodical><full-title>American Journal of Human Genetics</full-title><abbr-1>Am. J. Hum. Genet.</abbr-1><abbr-2>Am J Hum Genet</abbr-2></periodical><pages>1493-500</pages><volume>65</volume><dates><year>1999</year></dates><label>Broman1999</label><urls><related-urls><url><style face="underline" font="default" size="100%"> face="underline" font="default" size="100%">), Hutterites, villages in Sardinia and Friuli-Venezia-Giulia, ItalyPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5Fc2tvPC9BdXRob3I+PFllYXI+MjAxMzwvWWVhcj48UmVj
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ADDIN EN.CITE.DATA 19,38 reflecting serial bottlenecks during the dispersals across the globe. In these populations there are also often different levels of cryptic inbreeding indicated by long tails on the densities of SROH >10?Mb (e.g. for the Japanese shown in Fig. 2a, ~5% of the sample show evidence of recent inbreeding). Finally, a class of mixed populations present a heterogeneous picture, with admixed Native Americans and Hispanics/Latinos showing high variance in SROH, while African AmericansPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5QZW1iZXJ0b248L0F1dGhvcj48WWVhcj4yMDEyPC9ZZWFy
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ADDIN EN.CITE.DATA 46. The higher the Native American ancestry component, the greater the chance that these haplotypes will form ROH. The offspring of recent mixed race partnerships, on the other hand, will have very few ROH, given the low chance of shared parental haplotypes, irrespective of the particular continental ancestry.Socio-demographic factors that are not directly related to geography or principal components of ancestry can also influence ROH distributions. For example, in the Netherlands, which has a history of assortative mating by religious affiliation, FROH varies significantly between religious and non-religious groupsPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5BYmRlbGxhb3VpPC9BdXRob3I+PFllYXI+MjAxMzwvWWVh
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ADDIN EN.CITE PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5BYmRlbGxhb3VpPC9BdXRob3I+PFllYXI+MjAxMzwvWWVh
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ADDIN EN.CITE.DATA 47. Moreover, differential migration by educational status can induce systematic differences in FROH in highly educated, mobile versus less educated, less mobile population strataPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5BYmRlbGxhb3VpPC9BdXRob3I+PFllYXI+MjAxNTwvWWVh
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ADDIN EN.CITE.DATA 48. The effects of increased migration and urbanization through time also generate a secular trend in ROH such that younger European Americans have significantly lower burdens: NROH and SROH are predicted to have decreased by 14% and 24% across the last century ADDIN EN.CITE <EndNote><Cite><Author>Nalls</Author><Year>2009</Year><RecNum>102</RecNum><DisplayText><style face="superscript">49</style></DisplayText><record><rec-number>102</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487079810">102</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Nalls, Michael A</author><author>Simon-Sanchez, Javier</author><author>Gibbs, J Raphael</author><author>Paisan-Ruiz, Coro</author><author>Bras, Jose Tomas</author><author>Tanaka, Toshiko</author><author>Matarin, Mar</author><author>Scholz, Sonja</author><author>Weitz, Charles</author><author>Harris, Tamara B</author><author>Ferrucci, Luigi</author><author>Hardy, John</author><author>Singleton, Andrew B</author></authors></contributors><titles><title>Measures of autozygosity in decline: globalization, urbanization, and its implications for medical genetics</title><secondary-title>PLoS Genetics</secondary-title><short-title>Measures of autozygosity in decline: globalization, urbanization, and its implications for medical genetics</short-title></titles><periodical><full-title>PLoS Genetics</full-title><abbr-1>PLoS Genet.</abbr-1><abbr-2>PLoS Genet</abbr-2></periodical><pages>e1000415</pages><volume>5</volume><number>3</number><dates><year>2009</year></dates><isbn>1553-7404</isbn><accession-num>19282984</accession-num><label>eng</label><urls><related-urls><url> by Riadh Hammami</remote-database-provider></record></Cite></EndNote>49. [H3] Limits of homozygosity. Complete hydatidiform moles are a very rare form of non-viable pregnancy wherein the oocyte is enucleated and fertilized by a sperm. Thus the mole contains only sperm-derived DNA and is homozygous across the entire genome; they have been used to provide accurate haplotype mapsPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5IaWdhc2E8L0F1dGhvcj48WWVhcj4yMDA5PC9ZZWFyPjxS
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ADDIN EN.CITE.DATA 50. Uniparental disomy (UPD) occurs when both copies of a chromosome, or segment of a chromosome, are inherited from one parent, and therefore also generates ROH if two copies of one parental chromosome are present. However, the observation of Mendelian transmission of haplotypes giving rise to ROH demonstrates that most ROH are not due to UPD or other cytogenetic abnormalities ADDIN EN.CITE <EndNote><Cite><Author>Curtis</Author><Year>2007</Year><RecNum>42</RecNum><DisplayText><style face="superscript">34</style></DisplayText><record><rec-number>42</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487079810">42</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Curtis, David</author></authors></contributors><titles><title>Extended homozygosity is not usually due to cytogenetic abnormality</title><secondary-title>BMC Genetics</secondary-title><short-title>Extended homozygosity is not usually due to cytogenetic abnormality</short-title></titles><periodical><full-title>BMC Genetics</full-title></periodical><pages>67</pages><volume>8</volume><number>1</number><dates><year>2007</year></dates><isbn>1471-2156</isbn><label>Curtis2007</label><work-type>journal article</work-type><urls><related-urls><url>*~hmac=76cf92c3c2de703f9faef97b41abdc9365e74299c0f85b6ec6c1e1c37d185b22</url></related-urls></urls><electronic-resource-num>10.1186/1471-2156-8-67</electronic-resource-num></record></Cite></EndNote>34. Indeed, analysis of a large series of children with developmental delay or autism revealed UPD to be rare and to manifest with very long ROH, the shortest being 13.5?MbPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5QYXBlbmhhdXNlbjwvQXV0aG9yPjxZZWFyPjIwMTE8L1ll
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ADDIN EN.CITE.DATA 51. Thus UPD is unlikely to confound analyses of ROH to any great degree, particularly as subjects with ROH on multiple chromosomes can be excluded as UPD cases. Incest — mating between first-degree relatives — will generate an extreme burden of ROH, with ~25% of the genome expected to be in ROH. Several such cases have been found through clinical screening of children with intellectual disabilities or congenital abnormalities using microarraysPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5TY2hhYWY8L0F1dGhvcj48WWVhcj4yMDExPC9ZZWFyPjxS
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ADDIN EN.CITE.DATA 55 and the Ptolemies.In the data presented in Fig. 2, the most extreme individual has an SROH >500?Mb, including 342?Mb in ROH >10?Mb in length, probably the result of avuncular union [G] ADDIN EN.CITE <EndNote><Cite><Author>Leutenegger</Author><Year>2011</Year><RecNum>189</RecNum><DisplayText><style face="superscript">56</style></DisplayText><record><rec-number>189</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1488190516">189</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Leutenegger, A. L.</author><author>Sahbatou, M.</author><author>Gazal, S.</author><author>Cann, H.</author><author>Genin, E.</author></authors></contributors><auth-address>Inserm, U946, Paris, France. anne-louise.leutenegger@inserm.fr</auth-address><titles><title>Consanguinity around the world: what do the genomic data of the HGDP-CEPH diversity panel tell us?</title><secondary-title>European Journal of Human Genetics</secondary-title></titles><periodical><full-title>European Journal of Human Genetics</full-title><abbr-1>Europ. J. Hum. Genet.</abbr-1><abbr-2>Europ J Hum Genet</abbr-2></periodical><pages>583-7</pages><volume>19</volume><number>5</number><keywords><keyword>*Consanguinity</keyword><keyword>Female</keyword><keyword>Human Genome Project</keyword><keyword>Humans</keyword><keyword>Linkage Disequilibrium</keyword><keyword>Male</keyword><keyword>Marriage</keyword><keyword>*Models, Genetic</keyword><keyword>Pedigree</keyword></keywords><dates><year>2011</year><pub-dates><date>May</date></pub-dates></dates><isbn>1476-5438 (Electronic)
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ADDIN EN.CITE.DATA 35; only 5 of these had no ROH >10?Mb. Outside of populations with high mean SROH, first-cousin offspring are seen in two Japanese, one Uzbek, one South African Bantu-speaker, four Colombians and one Mexican (mestizos — recalling the Maracaibo Venezuelan family where ROH were first discovered ADDIN EN.CITE <EndNote><Cite><Author>Broman</Author><Year>1999</Year><RecNum>25</RecNum><DisplayText><style face="superscript">3</style></DisplayText><record><rec-number>25</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487079810">25</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Broman, K. W.</author><author>Weber, J. L.</author></authors></contributors><titles><title>Long homozygous chromosomal segments in reference families from the centre d'Etude du polymorphisme humain</title><secondary-title>American Journal of Human Genetics</secondary-title><short-title>Long homozygous chromosomal segments in reference families from the centre d'Etude du polymorphisme humain</short-title></titles><periodical><full-title>American Journal of Human Genetics</full-title><abbr-1>Am. J. Hum. Genet.</abbr-1><abbr-2>Am J Hum Genet</abbr-2></periodical><pages>1493-500</pages><volume>65</volume><dates><year>1999</year></dates><label>Broman1999</label><urls><related-urls><url><style face="underline" font="default" size="100%"> face="underline" font="default" size="100%">). Assessing FROH reveals not only interesting demographic historical information but allows prediction of the increased risk of rare recessive diseasesPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5KYWxraDwvQXV0aG9yPjxZZWFyPjIwMTU8L1llYXI+PFJl
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ADDIN EN.CITE.DATA 57.[H3] Distribution across the genome. ROH are somewhat more common in regions of high LD and low recombination ADDIN EN.CITE <EndNote><Cite><Author>Gibson</Author><Year>2006</Year><RecNum>55</RecNum><DisplayText><style face="superscript">29</style></DisplayText><record><rec-number>55</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487079810">55</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Gibson, J.</author><author>Morton, N. E.</author><author>Collins, A.</author></authors></contributors><titles><title>Extended tracts of homozygosity in outbred human populations</title><secondary-title>Human Molecular Genetics</secondary-title><short-title>Extended tracts of homozygosity in outbred human populations</short-title></titles><periodical><full-title>Human Molecular Genetics</full-title><abbr-1>Hum. Mol. Genet.</abbr-1><abbr-2>Hum Mol Genet</abbr-2></periodical><pages>789-795</pages><volume>15</volume><dates><year>2006</year></dates><label>Gibson2006</label><urls><related-urls><url><style face="underline" font="default" size="100%"> face="underline" font="default" size="100%">, are particularly prevalent on the X chromosomePEVuZE5vdGU+PENpdGU+PEF1dGhvcj5DdXJ0aXM8L0F1dGhvcj48WWVhcj4yMDA4PC9ZZWFyPjxS
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ADDIN EN.CITE.DATA 59. These observations are linked by low recombination: the X chromosome spends one third of its time in the male germline where (with the exception of the small pseudo-autosomal regions) it cannot recombine, and low recombination regions have low SNP diversity. Recombination breaks up chromosomal segments over generations and thus low recombination regions allow greater persistence of long ancestral haplotypes and an increased chance they come together to form ROH. Over and above this, there is a very uneven distribution along the genome, with a number of ‘ROH islands’ on each chromosome with significant excesses of ROHPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5MZW5jejwvQXV0aG9yPjxZZWFyPjIwMDc8L1llYXI+PFJl
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ADDIN EN.CITE.DATA 29,35. The common ancestors are recent enough that recombination has had little opportunity to break up the segments and so these are more randomly distributed across the genome. This difference is illustrated by the distribution of ROH >1?Mb in length on chromosome 1 (which reflects the genome-wide patternPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5DdXJ0aXM8L0F1dGhvcj48WWVhcj4yMDA4PC9ZZWFyPjxS
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ADDIN EN.CITE.DATA 58) in the relatively outbred Tuscans from the 1000 Genomes Project (Fig. 3a), versus that in the consanguineous Punjabis (Fig. 3b). There is a distinct tendency for Tuscans to carry comparatively short ROH in the same places — known as ROH islands — where commonly >10% of the population carries an ROH (Fig 3c). More randomly-sited ROH are also observed. Fine-scale investigation reveals remarkably consistent sharing of ROH boundaries from person-to-person, probably due to ancestral recombination eventsPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5Ob3RobmFnZWw8L0F1dGhvcj48WWVhcj4yMDEwPC9ZZWFy
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ADDIN EN.CITE.DATA 59 (and once more highlighting the pervasive influence of recombination on ROH distributions). Whereas ROH islands are also present in the Punjabis, in some cases at the same loci as in Europeans, a significant minority of the population carries much longer ROH scattered across the genome, elevating the baseline proportion of individuals who are autozygous (Fig. 3d). In some cases ROH islands are due to homozygosity of one common haplotype but in other cases, multiple different haplotypes contribute to the ROHPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5DdXJ0aXM8L0F1dGhvcj48WWVhcj4yMDA4PC9ZZWFyPjxS
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ADDIN EN.CITE.DATA 58. The origin of ROH islands is subject to debate, but it appears that there are extended haplotypes segregating at high frequencies in the population in these regions. In some cases this can be explained by selection; for example, there is an ROH island around the lactase (LCT) gene on chromosome 2q21 in Europeans, the site of very strong selection for the ability to metabolise lactose as an adultPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5DdXJ0aXM8L0F1dGhvcj48WWVhcj4yMDA4PC9ZZWFyPjxS
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ADDIN EN.CITE.DATA 58, and numerous other islands are probable targets of recent positive selectionPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5QZW1iZXJ0b248L0F1dGhvcj48WWVhcj4yMDEyPC9ZZWFy
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ADDIN EN.CITE.DATA 58 — or some may not be truly autozygous. Whole-genome sequence data will facilitate an assessment of whether hitherto ungenotyped variants in ROH islands are also homozygous and thus the potential contribution of rare variants in these regions to disease risk. [H1] Runs of homozygosity and complex diseaseAlthough homozygosity mapping ADDIN EN.CITE <EndNote><Cite><Author>Lander</Author><Year>1987</Year><RecNum>84</RecNum><DisplayText><style face="superscript">60</style></DisplayText><record><rec-number>84</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487079810">84</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Lander, E. S.</author><author>Botstein, D.</author></authors></contributors><titles><title>Homozygosity mapping - A way to map human recessive traits with the DNA of inbred children</title><secondary-title>Science</secondary-title><short-title>HOMOZYGOSITY MAPPING - A WAY TO MAP HUMAN RECESSIVE TRAITS WITH THE DNA OF INBRED CHILDREN</short-title></titles><periodical><full-title>Science</full-title></periodical><pages>1567-1570</pages><volume>236</volume><number>4808</number><dates><year>1987</year><pub-dates><date>Jun</date></pub-dates></dates><isbn>0036-8075</isbn><accession-num>WOS:A1987H781800028</accession-num><urls><related-urls><url><Go to ISI>://WOS:A1987H781800028</url><url> has successfully identified the loci underlying many hundreds of rare recessive disorders, mostly in high-homozygosity populations, attention has only recently turned to the relationship between ROH and complex diseases ADDIN EN.CITE <EndNote><Cite><Author>Rudan</Author><Year>2006</Year><RecNum>190</RecNum><DisplayText><style face="superscript">61</style></DisplayText><record><rec-number>190</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1488190637">190</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Rudan, I.</author><author>Campbell, H.</author><author>Carothers, A. D.</author><author>Hastie, N. D.</author><author>Wright, A. F.</author></authors></contributors><titles><title>Contribution of consanguinuity to polygenic and multifactorial diseases</title><secondary-title>Nature Genetics</secondary-title></titles><periodical><full-title>Nature Genetics</full-title><abbr-1>Nat. Genet.</abbr-1><abbr-2>Nat Genet</abbr-2></periodical><pages>1224-5</pages><volume>38</volume><number>11</number><keywords><keyword>Animals</keyword><keyword>*Consanguinity</keyword><keyword>Genetic Diseases, Inborn/*epidemiology/*etiology</keyword><keyword>Humans</keyword><keyword>Middle East/epidemiology</keyword><keyword>Multifactorial Inheritance/*genetics</keyword></keywords><dates><year>2006</year><pub-dates><date>Nov</date></pub-dates></dates><isbn>1061-4036 (Print)
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ADDIN EN.CITE.DATA 7. Power also depends on the variance in SROH, which is highly correlated with mean SROH, such that more homozygous populations are more powerful. An interesting example is provided by serial analyses of schizophrenia risk and ROH, whereby an initial meta-analysis of >9,000 cases and >12,000 controls provided evidence that a 1% increase in FROH conferred a 17% increase in risk of schizophreniaPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5LZWxsZXI8L0F1dGhvcj48WWVhcj4yMDEyPC9ZZWFyPjxS
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ADDIN EN.CITE.DATA 62. However, analysis of a much larger sample set, totaling nearly 40,000 subjects found a much attenuated signal and concluded there was no reliable association between burden of ROH and case statusPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5Kb2huc29uPC9BdXRob3I+PFllYXI+MjAxNjwvWWVhcj48
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ADDIN EN.CITE.DATA 63. Confounding [G] , publication bias and other biases may therefore also be at play ADDIN EN.CITE <EndNote><Cite><Author>Panagiotou</Author><Year>2013</Year><RecNum>220</RecNum><DisplayText><style face="superscript">64</style></DisplayText><record><rec-number>220</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1489047231">220</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Panagiotou, O. A.</author><author>Willer, C. J.</author><author>Hirschhorn, J. N.</author><author>Ioannidis, J. P.</author></authors></contributors><auth-address>Clinical and Molecular Epidemiology Unit, Department of Hygiene and Epidemiology, University of Ioannina School of Medicine, Ioannina 45110, Greece; email: orestis.panagiotou@.</auth-address><titles><title>The power of meta-analysis in genome-wide association studies</title><secondary-title>The Annual Review of Genomics and Human Genetics</secondary-title></titles><periodical><full-title>The Annual Review of Genomics and Human Genetics</full-title><abbr-1>Annu. Rev. Genomics. Hum. Genet.</abbr-1></periodical><pages>441-65</pages><volume>14</volume><keywords><keyword>Genetics, Medical</keyword><keyword>*Genome, Human</keyword><keyword>*Genome-Wide Association Study</keyword><keyword>HapMap Project</keyword><keyword>Humans</keyword><keyword>Meta-Analysis as Topic</keyword></keywords><dates><year>2013</year></dates><isbn>1545-293X (Electronic)
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ADDIN EN.CITE.DATA 47. Further complications may arise due to ascertainment biases between cases and controls, or experimental biases, e.g. genotyping and ROH calling not being performed jointly for cases and controls. For ROH, this may be particularly sensitive as the precise length, and evidence for the existence of an ROH may depend on the density (and of course accuracy) of the genotyping microarray usedbXBlcmlhbCBDb2xsZWdlIExvbmRvbiwgTm9yZm9sayBQbGFjZSwgTG9uZG9uIFcyIDFQRywgVUsu
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ADDIN EN.CITE.DATA 20.[H3] Quantitative traits. More consistency has been observed in studies of the association between SROH and quantitative traits, perhaps due to the larger sample sizes, the use of common microarrays within study populations, and the avoidance of unmatched controls. An exceptionally large study of up to 354,224 subjects found βFROH, the regression coefficient between trait and the proportion of the genome in ROH, to be –2.9 (0.2), –3.5 (0.7), –4.7 (0.6) and –4.6 (0.7) phenotypic standard deviations (standard errors in brackets) for height, forced expiratory lung volume in one second, cognitive ability and educational attainmentbXBlcmlhbCBDb2xsZWdlIExvbmRvbiwgTm9yZm9sayBQbGFjZSwgTG9uZG9uIFcyIDFQRywgVUsu
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ADDIN EN.CITE.DATA 20 (Fig. 4). The offspring of first cousins are thus predicted to be 1.2 cm shorter, have 140 ml lower lung expiratory capacity, 0.29 standard deviations less generalized cognitive function (~4.3 IQ points) and attain 9.7 fewer months of education. Although the association with height was already knownPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5NY1F1aWxsYW48L0F1dGhvcj48WWVhcj4yMDEyPC9ZZWFy
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ADDIN EN.CITE.DATA 67 and effect sizes were consistent with pedigree-based estimates for stature and IQ ADDIN EN.CITE <EndNote><Cite><Author>Morton</Author><Year>1978</Year><RecNum>212</RecNum><DisplayText><style face="superscript">68,69</style></DisplayText><record><rec-number>212</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1488618900">212</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Morton, N. E.</author></authors></contributors><titles><title>Effects of inbreeding on IQ and mental retardation</title><secondary-title>Proceedings of the National Academy of Sciences of the United States of America</secondary-title></titles><periodical><full-title>Proceedings of the National Academy of Sciences of the United States of America</full-title><abbr-1>Proc. Natl. Acad. Sci. U. S. A.</abbr-1><abbr-2>Proc Natl Acad Sci U S A</abbr-2></periodical><pages>3906-3908</pages><volume>75</volume><dates><year>1978</year></dates><urls></urls></record></Cite><Cite><Author>Schull</Author><Year>1962</Year><RecNum>217</RecNum><record><rec-number>217</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1488702825">217</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Schull, W. J.</author></authors></contributors><titles><title>Inbreeding and maternal effects in the Japanese</title><secondary-title>Eugenetics Quaterly</secondary-title></titles><periodical><full-title>Eugenetics Quaterly</full-title><abbr-1>Eugen. Q.</abbr-1></periodical><pages>14-22</pages><volume>9</volume><dates><year>1962</year></dates><urls></urls></record></Cite></EndNote>68,69. However, no effects of homozygosity were observed for a series of cardiometabolic risk factors, despite very large sample sizesbXBlcmlhbCBDb2xsZWdlIExvbmRvbiwgTm9yZm9sayBQbGFjZSwgTG9uZG9uIFcyIDFQRywgVUsu
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ADDIN EN.CITE.DATA 20.Despite consistency across these studies, it remains important to consider confounding. One study of ~2,000 subjects of Dutch ancestry showed that parental educational attainment is very strongly related to offspring SROH, a relationship fully explained by the distance between the parents’ birthplaces, i.e. more educated individuals moved further before finding a spouse, and the two parents were thus, on average, less genomically relatedPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5BYmRlbGxhb3VpPC9BdXRob3I+PFllYXI+MjAxNTwvWWVh
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ADDIN EN.CITE.DATA 20 also showed that the effect sizes they observed were similar in populations with higher and lower mean SROH, a consistency expected if the effect is from SROH, but not expected under a model of confounding by socio-economic status. The signals of association were also robust to population stratification [G] by geography or demographic history, and inclusion of educational attainment as a covariatebXBlcmlhbCBDb2xsZWdlIExvbmRvbiwgTm9yZm9sayBQbGFjZSwgTG9uZG9uIFcyIDFQRywgVUsu
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ADDIN EN.CITE.DATA 20. As expected under the directional dominance model, associations were also found in populations for which there is very limited migration and sometimes communal living, such as the Hutterites, Amish and Italian hill towns and mountain villages. Fitness-related traits are thought to show inbreeding depression and/or directional dominance, due to the limited effectiveness of purifying selection on rare, deleterious recessive alleles. The associations between SROH and stature and cognition thus suggest that these traits — or perhaps more likely, some underlying trait or traits — are components of Darwinian fitness [G] and have a rare, recessive element to their genetic architecture ADDIN EN.CITE <EndNote><Cite><Author>Charlesworth</Author><Year>2009</Year><RecNum>36</RecNum><DisplayText><style face="superscript">13</style></DisplayText><record><rec-number>36</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487079810">36</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Charlesworth, D.</author><author>Willis, J. H.</author></authors></contributors><titles><title>The genetics of inbreeding depression</title><secondary-title>Nature Reviews Genetics</secondary-title><short-title>FUNDAMENTAL CONCEPTS IN GENETICS The genetics of inbreeding depression</short-title></titles><periodical><full-title>Nature Reviews Genetics</full-title><abbr-1>Nat. Rev. Genet.</abbr-1><abbr-2>Nat Rev Genet</abbr-2></periodical><pages>783-796</pages><volume>10</volume><number>11</number><dates><year>2009</year><pub-dates><date>Nov</date></pub-dates></dates><isbn>1471-0056</isbn><accession-num>WOS:000270915400014</accession-num><urls><related-urls><url><Go to ISI>://WOS:000270915400014</url></related-urls></urls><electronic-resource-num>10.1038/nrg2664</electronic-resource-num></record></Cite></EndNote>13. The contribution of this component to genetic variance may be low, should the individual alleles be very rare and fully recessive, even if there are many such alleles. However, the decline in ROH due to increasing panmixia [G] and urbanization will be beneficial in terms of reducing the burden of recessive Mendelian disease and risk factors for complex traits showing directional dominance ADDIN EN.CITE <EndNote><Cite><Author>Rudan</Author><Year>2008</Year><RecNum>216</RecNum><DisplayText><style face="superscript">70</style></DisplayText><record><rec-number>216</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1488702628">216</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Rudan, I.</author><author>Carothers, A. D.</author><author>Polasek, O.</author><author>Hayward, C.</author><author>Vitart, V.</author><author>Biloglav, Z.</author><author>Kolcic, I.</author><author>Zgaga, L.</author><author>Ivankovic, D.</author><author>Vorko-Jovic, A.</author><author>Wilson, J. F.</author><author>Weber, J. L.</author><author>Hastie, N.</author><author>Wright, A.</author><author>Campbell, H.</author></authors></contributors><auth-address>Department of Public Health Sciences, University of Edinburgh, Edinburgh, UK.</auth-address><titles><title>Quantifying the increase in average human heterozygosity due to urbanisation</title><secondary-title>European Journal of Human Genetics</secondary-title></titles><periodical><full-title>European Journal of Human Genetics</full-title><abbr-1>Europ. J. Hum. Genet.</abbr-1><abbr-2>Europ J Hum Genet</abbr-2></periodical><pages>1097-102</pages><volume>16</volume><number>9</number><keywords><keyword>Consanguinity</keyword><keyword>Croatia</keyword><keyword>Genetic Markers</keyword><keyword>Genetic Variation</keyword><keyword>Genetics, Population</keyword><keyword>*Heterozygote Detection</keyword><keyword>Humans</keyword><keyword>Rural Population</keyword><keyword>Urban Population</keyword><keyword>*Urbanization</keyword></keywords><dates><year>2008</year><pub-dates><date>Sep</date></pub-dates></dates><isbn>1018-4813 (Print)
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ADDIN EN.CITE.DATA 72, these associations are not robust and have not been replicated. Thus although homozygosity mapping in inbred populations has been exceptionally successful for monogenic recessive disorders, ROH mapping studies have had less success for complex traits. Nevertheless, when performed in well-powered samples such as the UK Biobank and other very large studies, individual ROH associations have the advantage of potentially being able to identify rare recessive variants cost efficiently, without large-scale WGS.[H1] Final considerationsROH studies are proliferating, with applications to diverse topics, from enhanced discovery of human knockouts to estimation of the human mutation rate (Box 3). Isolated populations yield a higher number of homozygous loss-of-function mutations per sequence compared to cosmopolitan groupsPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5LYWlzZXI8L0F1dGhvcj48WWVhcj4yMDE1PC9ZZWFyPjxS
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RW5kTm90ZT4A
ADDIN EN.CITE.DATA 54. In some circumstances it is likely that these differences could not only influence measured ROH, but also the apparent strength of association with traits. To help aid comparability of results, we suggest that studies of ROH using microarray data should adopt the following criteria. The microarray should be genome-wide, have at least 300,000 SNPs and results presented should include βFROH (accompanied by standard error) using the same protocol as the ROHgen consortiumbXBlcmlhbCBDb2xsZWdlIExvbmRvbiwgTm9yZm9sayBQbGFjZSwgTG9uZG9uIFcyIDFQRywgVUsu
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ADDIN EN.CITE.DATA 20: a run of homozygosity, called by PLINK should consider SNPs >5% allele frequency, have at least 50 SNPs and be 1.5?Mb long, with allowance for missing and heterozygous calls. This is not to preclude presentation of central results on another basis, for example also including shorter ROH, but presentation somewhere against a common baseline would enable readers to compare and contrast results, and see how changes in method influence results. The inbreeding coefficient FROH can be calculated as the SROH divided by the length of the autosomal genome. The continued evolution of sequencing technologies means more research is required before recommendations can be drafted for ROH calling with these data. Good practices in quantitative genetics such as large sample sizes, control of population structure, replication studies and accurate matching of cases and controls, or better using case–cohort studies, will increase the likelihood of robust findings.[H3] Future directions. The very large genomic datasets now becoming availablePEVuZE5vdGU+PENpdGU+PEF1dGhvcj5TdWRsb3c8L0F1dGhvcj48WWVhcj4yMDE1PC9ZZWFyPjxS
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ADDIN EN.CITE.DATA 76-79 offer researchers a unique opportunity to better understand the influence of ROH on complex disease architecture. Such datasets will allow well-powered, broad surveys of phenotypes — including omic analytes that are mechanistically proximal to the gene and potentially fitness-related immune traits — to delineate the scale of inbreeding depression and to identify genomic regions with recessive effects on both complex traits and rare Mendelian diseases. A number of future research questions are suggested here. Does the burden of homozygosity caused by reduced population size have the same effect as recent consanguinity and can this be used to infer whether the variants responsible for inbreeding depression are rare or common, recessive or over-dominant? Recent consanguinity gives rise to long ROH which bring almost all variants, from common to very rare, into a homozygous state. By contrast, more distant inbreeding causes shorter ROH which only make homozygous the variants present in the shared ancestor, which are by definition common variants. If (unobserved) mutations have occurred in either the maternal or paternal line since the time of a common ancestor, these will not be homozygous in the inbred individual. Thus, analysing the effect of different lengths of ROH may reveal the relative contributions of rare and common variants: greater effects per megabase for larger ROH imply that rarer variants are causing the inbreeding depression.Is there any evidence that mixed-race individuals differ in fitness-related traits from their peers, either due to heterosis [G] or outbreeding depression [G] ? Is the effect of inbreeding sex-specific in humans, as has been observed in other species ADDIN EN.CITE <EndNote><Cite><Author>Ebel</Author><Year>2016</Year><RecNum>204</RecNum><DisplayText><style face="superscript">80</style></DisplayText><record><rec-number>204</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1488575343">204</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Ebel, E. R.</author><author>Phillips, P. C.</author></authors></contributors><auth-address>Institute of Ecology and Evolution and Department of Biology, 5289 University of Oregon, 97403, Eugene, Oregon, USA.
Present address: Department of Biology, Stanford University, 371 Serra Mall, Stanford, CA, 94305, USA.
Institute of Ecology and Evolution and Department of Biology, 5289 University of Oregon, 97403, Eugene, Oregon, USA. pphil@uoregon.edu.</auth-address><titles><title>Intrinsic differences between males and females determine sex-specific consequences of inbreeding</title><secondary-title>BMC Evolutionary Biology</secondary-title></titles><periodical><full-title>BMC Evolutionary Biology</full-title><abbr-1>BMC Evol. Biol.</abbr-1><abbr-2>BMC Evol Biol</abbr-2></periodical><pages>36</pages><volume>16</volume><keywords><keyword>Animals</keyword><keyword>Biological Evolution</keyword><keyword>Caenorhabditis/*genetics/physiology</keyword><keyword>Female</keyword><keyword>Homozygote</keyword><keyword>*Inbreeding</keyword><keyword>Male</keyword><keyword>Mating Preference, Animal</keyword><keyword>Mutation</keyword><keyword>Reproduction/genetics/physiology</keyword><keyword>*Sex Characteristics</keyword><keyword>Sex Factors</keyword></keywords><dates><year>2016</year><pub-dates><date>Feb 09</date></pub-dates></dates><isbn>1471-2148 (Electronic)
1471-2148 (Linking)</isbn><accession-num>26860745</accession-num><urls><related-urls><url>? Is it possible to identify specific genomic regions where ROH influence complex traits, and if so, do these loci correspond to known GWAS hits, or does regional ROH mapping offer a complementary method for identifying novel biology? One of the strengths of ROH analyses is that long homozygous segments can be reliably identified even from relatively modest marker densities. However, the increasing availability of WGS will soon allow shorter ROH to be more reliably called, in larger datasets, than is currently possible. This should permit the effect of very short ROH on disease risk to be quantified, as well as potentially shedding further light on the demographic history of human populations. ROH studies will further illuminate the scope and mechanism of inbreeding depression in humans.BoxesBox 1 | Inbreeding depression, Charles Darwin and Royal Dynasties.The first research programme on the harmful effects of inbreeding, the mating of close relatives, was performed by Charles Darwin ADDIN EN.CITE <EndNote><Cite><Author>Darwin</Author><Year>1868</Year><RecNum>166</RecNum><DisplayText><style face="superscript">81,82</style></DisplayText><record><rec-number>166</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487243492">166</key></foreign-keys><ref-type name="Book">6</ref-type><contributors><authors><author>Darwin, C. R.</author></authors></contributors><titles><title>The Variation of animals and plants under domestication</title></titles><dates><year>1868</year></dates><pub-location>London</pub-location><publisher>John Murray</publisher><urls></urls></record></Cite><Cite><Author>Darwin</Author><Year>1876</Year><RecNum>167</RecNum><record><rec-number>167</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487243637">167</key></foreign-keys><ref-type name="Book">6</ref-type><contributors><authors><author>Darwin, C. R.</author></authors></contributors><titles><title>The effects of cross and self fertilisation in the vegetable kingdom</title></titles><dates><year>1876</year></dates><pub-location>London</pub-location><publisher>John Murray</publisher><urls></urls></record></Cite></EndNote>81,82. He carried out carefully controlled experiments in plants that involved self-fertilization and outcrossing between unrelated individuals in 57 species and showed that the offspring of self-fertilized plants were on average shorter, flowered later and produced fewer seeds than the progeny of cross-fertilized plants. He thus documented the phenomenon of inbreeding depression, the decline of traits that are closely related to fitness, now known to be caused by the increase in homozygosity in inbred individuals. Darwin also had a personal interest in the adverse effects of inbreeding since he was married to his first cousin Emma Wedgwood (see the figure), and they had ten children who were often ill, three of whom died at an early age ADDIN EN.CITE <EndNote><Cite><Author>Berra</Author><Year>2013</Year><RecNum>168</RecNum><DisplayText><style face="superscript">83</style></DisplayText><record><rec-number>168</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487243882">168</key></foreign-keys><ref-type name="Book">6</ref-type><contributors><authors><author>Berra, T. M.</author></authors></contributors><titles><title>Darwin & his children: his other legacy</title></titles><dates><year>2013</year></dates><pub-location>New York</pub-location><publisher>Oxford University Press</publisher><urls></urls></record></Cite></EndNote>83. Charles and Emma will each have inherited large segments of the genomes of their grandparents Josiah Wedgwood I and Sarah Wedgwood identical-by-descent, and transmitted some of these to their children, thus generating long runs of homozygosity (ROH) wherever the same segments have been passed down each side of the pedigree. Darwin’s concerns about the harmful effects of first-cousin marriage in his progeny have been considered exaggerated because they were based on the extrapolation from ill-effects of self-fertilization in plants to the outcomes of first-cousin marriage in humans. However, the possibility of inbreeding effects on the Darwin children is supported by the decrease in both childhood survival and male fertility detected in the progeny of a number of consanguineous marriages of the Darwin-Wedgwood dynastyPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5CZXJyYTwvQXV0aG9yPjxZZWFyPjIwMTA8L1llYXI+PFJl
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ADDIN EN.CITE.DATA 84,85.Although studies in the Hutterites have shown a decrease in fecundity for inbred women (as well as evidence of reproductive compensation) ADDIN EN.CITE <EndNote><Cite><Author>Ober</Author><Year>1999</Year><RecNum>175</RecNum><DisplayText><style face="superscript">86</style></DisplayText><record><rec-number>175</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487923270">175</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Ober, C.</author><author>Hyslop, T.</author><author>Hauck, W. W.</author></authors></contributors><auth-address>Departments of Human Genetics and Obstetrics and Gynecology, The University of Chicago, IL 60637, USA. carole@genetics.uchicago.edu</auth-address><titles><title>Inbreeding effects on fertility in humans: evidence for reproductive compensation</title><secondary-title>American Journal of Human Genetics</secondary-title></titles><periodical><full-title>American Journal of Human Genetics</full-title><abbr-1>Am. J. Hum. Genet.</abbr-1><abbr-2>Am J Hum Genet</abbr-2></periodical><pages>225-31</pages><volume>64</volume><number>1</number><keywords><keyword>Adult</keyword><keyword>*Consanguinity</keyword><keyword>Female</keyword><keyword>Fertility/*genetics</keyword><keyword>Humans</keyword><keyword>Male</keyword><keyword>Maternal Age</keyword><keyword>Parity</keyword><keyword>Pregnancy</keyword><keyword>Pregnancy Outcome</keyword><keyword>Survival Analysis</keyword></keywords><dates><year>1999</year><pub-dates><date>Jan</date></pub-dates></dates><isbn>0002-9297 (Print)
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ADDIN EN.CITE.DATA 88-90. In the House of Habsburg, strong inbreeding depression for both infant and child mortality was detected c.1450–1800. A considerable reduction of this inbreeding effect on child survival in a fairly small number of generations was observed, potentially caused by the purging of deleterious alleles of a large effect, a mechanism previously observed for loss-of-function alleles in mountain gorillasPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5YdWU8L0F1dGhvcj48WWVhcj4yMDE1PC9ZZWFyPjxSZWNO
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ADDIN EN.CITE.DATA 16.Box 2 | ROH in ancient humans, Neanderthals and great apes.The ability to generate genome-wide genotypes or whole genome sequences from ancient DNA has ushered in a new era in understanding human population history, including via runs of homozygosity (ROH). It is striking that both Upper Palaeolithic and Mesolithic hunter-gatherers, from Luxembourg, Switzerland and Georgia (~6000–11,000 BC), carried very high levels of ROH, comparable to modern Oceanians and certain Siberian, Indian and Greenlander populations. By contrast Neolithic skeletons from Northern Ireland, Hungary, South-West Germany and Anatolia (~3200–6000 BC), showed much reduced levels, comparable to modern East Asians, with Bronze Age samples from Northern Ireland and Hungary (~2000–1200 BC) even lower, similar to modern EuropeansPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5DYXNzaWR5PC9BdXRob3I+PFllYXI+MjAxNjwvWWVhcj48
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ADDIN EN.CITE.DATA 42,91-93 (see the figure). Plotted in the main figure ADDIN EN.CITE <EndNote><Cite><Author>Cassidy</Author><Year>2016</Year><RecNum>32</RecNum><DisplayText><style face="superscript">91</style></DisplayText><record><rec-number>32</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487079810">32</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Cassidy, Lara M</author><author>Martiniano, Rui</author><author>Murphy, Eileen M</author><author>Teasdale, Matthew D</author><author>Mallory, James</author><author>Hartwell, Barrie</author><author>Bradley, Daniel G</author></authors></contributors><titles><title>Neolithic and Bronze Age migration to Ireland and establishment of the insular Atlantic genome</title><secondary-title>Proceedings of the National Academy of Sciences of the United States of America</secondary-title><short-title>Neolithic and Bronze Age migration to Ireland and establishment of the insular Atlantic genome</short-title></titles><periodical><full-title>Proceedings of the National Academy of Sciences of the United States of America</full-title><abbr-1>Proc. Natl. Acad. Sci. U. S. A.</abbr-1><abbr-2>Proc Natl Acad Sci U S A</abbr-2></periodical><pages>368-73</pages><volume>113</volume><number>2</number><dates><year>2016</year></dates><isbn>1091-6490</isbn><accession-num>26712024</accession-num><label>eng</label><urls><related-urls><url><style face="underline" font="default" size="100%"> face="underline" font="default" size="100%"> by Riadh Hammami</remote-database-provider></record></Cite></EndNote>91 is the sum of ROH in different megabase (Mb) length categories for one Mesolithic (bold black line), three Neolithic (bold blue lines) and two Bronze Age (bold red lines) samples, along with representative individuals from modern populations (thin lines with colours indicating continent of origin). The results imply that the Western and Caucasus hunter-gatherers lived in relatively isolated, endogamous societies, unlike the Neolithic farmers who appear to have arrived as part of a folk migration with a large effective population size. There is in fact a relationship between the median calibrated carbon-14 date of Hungarian remains and sum total length of ROH (SROH) (r2=0.4, p=0.06) ADDIN EN.CITE <EndNote><Cite><Author>Gamba</Author><Year>2014</Year><RecNum>50</RecNum><DisplayText><style face="superscript">92</style></DisplayText><record><rec-number>50</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487079810">50</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Gamba, Cristina</author><author>Jones, Eppie R</author><author>Teasdale, Matthew D</author><author>McLaughlin, Russell L</author><author>Gonzalez-Fortes, Gloria</author><author>Mattiangeli, Valeria</author><author>Domboróczki, László</author><author>K?vári, Ivett</author><author>Pap, Ildikó</author><author>Anders, Alexandra</author><author>Whittle, Alasdair</author><author>Dani, János</author><author>Raczky, Pál</author><author>Higham, Thomas F G</author><author>Hofreiter, Michael</author><author>Bradley, Daniel G</author><author>Pinhasi, Ron</author></authors></contributors><titles><title>Genome flux and stasis in a five millennium transect of European prehistory</title><secondary-title>Nature Communications</secondary-title><short-title>Genome flux and stasis in a five millennium transect of European prehistory</short-title></titles><periodical><full-title>Nature Communications</full-title><abbr-1>Nat. Commun.</abbr-1></periodical><pages>5257</pages><volume>5</volume><dates><year>2014</year></dates><isbn>2041-1723</isbn><accession-num>25334030</accession-num><label>eng</label><urls><related-urls><url> by Riadh Hammami</remote-database-provider></record></Cite></EndNote>92 (see the figure inset). Plotted in the figure inset ADDIN EN.CITE <EndNote><Cite><Author>Gamba</Author><Year>2014</Year><RecNum>50</RecNum><DisplayText><style face="superscript">92</style></DisplayText><record><rec-number>50</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487079810">50</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Gamba, Cristina</author><author>Jones, Eppie R</author><author>Teasdale, Matthew D</author><author>McLaughlin, Russell L</author><author>Gonzalez-Fortes, Gloria</author><author>Mattiangeli, Valeria</author><author>Domboróczki, László</author><author>K?vári, Ivett</author><author>Pap, Ildikó</author><author>Anders, Alexandra</author><author>Whittle, Alasdair</author><author>Dani, János</author><author>Raczky, Pál</author><author>Higham, Thomas F G</author><author>Hofreiter, Michael</author><author>Bradley, Daniel G</author><author>Pinhasi, Ron</author></authors></contributors><titles><title>Genome flux and stasis in a five millennium transect of European prehistory</title><secondary-title>Nature Communications</secondary-title><short-title>Genome flux and stasis in a five millennium transect of European prehistory</short-title></titles><periodical><full-title>Nature Communications</full-title><abbr-1>Nat. Commun.</abbr-1></periodical><pages>5257</pages><volume>5</volume><dates><year>2014</year></dates><isbn>2041-1723</isbn><accession-num>25334030</accession-num><label>eng</label><urls><related-urls><url> by Riadh Hammami</remote-database-provider></record></Cite></EndNote>92 is the correlation between median calibrated dates before the common era (BC, in thousands of years (ky)) and SROH for 9 ancient Hungarian skeletons. Additionally, the very early (~108,000 BC) Neolithic sample from Boncuklu in Anatolia, who was probably an indigenous forager that adopted cultivation, was intermediate in ROH distribution to the Mesolithic and later Neolithic samplesPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5LxLFsxLFuw6c8L0F1dGhvcj48WWVhcj4yMDE2PC9ZZWFy
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ADDIN EN.CITE.DATA 95.The distribution of ROH in hominins can be put into perspective by comparing to the other great apes. The endangered mountain gorillas have exceptionally high levels of homozygosity, with an average of 34% of their genomes in ROHPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5YdWU8L0F1dGhvcj48WWVhcj4yMDE1PC9ZZWFyPjxSZWNO
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ADDIN EN.CITE.DATA 16. 19% of their genomes consist of ROH between 2.5–10?Mb, easily more than the most homozygous reported human, and the Altai Neanderthal. The homozygosity implies several generations of recent as well as ancient inbreeding in the ancestry of the seven individuals sampled. Eastern lowland gorillas also show exceedingly high levels of ROH, about double the sum and number of ROH that are typical among the KaritianaPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5QcmFkby1NYXJ0aW5lejwvQXV0aG9yPjxZZWFyPjIwMTM8
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ADDIN EN.CITE.DATA 96. By contrast, Western lowland gorillas, chimpanzee, bonobo and orang-utan subspecies are much less homozygous, even if often considerably higher than most outbred human populations, averaging Oceanian levels for bonobos, for instance. Thus the great majority of humans are at the lower end of the hominid homozygosity spectrum, and only very isolated populations reflect the pattern that is prevalent in most great apes, where habitat fragmentation has reduced breeding population sizes dramatically. Eastern lowland and mountain gorillas are considerably more homozygous than any human or other great ape population.The figure is adapted from Refs ADDIN EN.CITE <EndNote><Cite><Author>Cassidy</Author><Year>2016</Year><RecNum>32</RecNum><DisplayText><style face="superscript">91</style></DisplayText><record><rec-number>32</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487079810">32</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Cassidy, Lara M</author><author>Martiniano, Rui</author><author>Murphy, Eileen M</author><author>Teasdale, Matthew D</author><author>Mallory, James</author><author>Hartwell, Barrie</author><author>Bradley, Daniel G</author></authors></contributors><titles><title>Neolithic and Bronze Age migration to Ireland and establishment of the insular Atlantic genome</title><secondary-title>Proceedings of the National Academy of Sciences of the United States of America</secondary-title><short-title>Neolithic and Bronze Age migration to Ireland and establishment of the insular Atlantic genome</short-title></titles><periodical><full-title>Proceedings of the National Academy of Sciences of the United States of America</full-title><abbr-1>Proc. Natl. Acad. Sci. U. S. A.</abbr-1><abbr-2>Proc Natl Acad Sci U S A</abbr-2></periodical><pages>368-73</pages><volume>113</volume><number>2</number><dates><year>2016</year></dates><isbn>1091-6490</isbn><accession-num>26712024</accession-num><label>eng</label><urls><related-urls><url><style face="underline" font="default" size="100%"> face="underline" font="default" size="100%"> by Riadh Hammami</remote-database-provider></record></Cite></EndNote>91, ADDIN EN.CITE <EndNote><Cite><Author>Gamba</Author><Year>2014</Year><RecNum>50</RecNum><DisplayText><style face="superscript">92</style></DisplayText><record><rec-number>50</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487079810">50</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Gamba, Cristina</author><author>Jones, Eppie R</author><author>Teasdale, Matthew D</author><author>McLaughlin, Russell L</author><author>Gonzalez-Fortes, Gloria</author><author>Mattiangeli, Valeria</author><author>Domboróczki, László</author><author>K?vári, Ivett</author><author>Pap, Ildikó</author><author>Anders, Alexandra</author><author>Whittle, Alasdair</author><author>Dani, János</author><author>Raczky, Pál</author><author>Higham, Thomas F G</author><author>Hofreiter, Michael</author><author>Bradley, Daniel G</author><author>Pinhasi, Ron</author></authors></contributors><titles><title>Genome flux and stasis in a five millennium transect of European prehistory</title><secondary-title>Nature Communications</secondary-title><short-title>Genome flux and stasis in a five millennium transect of European prehistory</short-title></titles><periodical><full-title>Nature Communications</full-title><abbr-1>Nat. Commun.</abbr-1></periodical><pages>5257</pages><volume>5</volume><dates><year>2014</year></dates><isbn>2041-1723</isbn><accession-num>25334030</accession-num><label>eng</label><urls><related-urls><url> by Riadh Hammami</remote-database-provider></record></Cite></EndNote>92.Box 3 | Estimating the human mutation rate from ROH. The mutation rate is a central parameter in biology — being the key to timing of the molecular clock as well as informing our expectations of mutations in cancer and understanding the incidence of genetic diseases — yet it has proved remarkably difficult to ascertain accurately ADDIN EN.CITE <EndNote><Cite><Author>Callaway</Author><Year>2015</Year><RecNum>30</RecNum><DisplayText><style face="superscript">97</style></DisplayText><record><rec-number>30</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487079810">30</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Callaway, E.</author></authors></contributors><titles><title>DNA mutation clock proves tough to set</title><secondary-title>Nature</secondary-title><short-title>BIOLOGY DNA clock proves tough to set</short-title></titles><periodical><full-title>Nature</full-title></periodical><pages>139-140</pages><volume>519</volume><number>7542</number><dates><year>2015</year><pub-dates><date>Mar</date></pub-dates></dates><isbn>0028-0836</isbn><accession-num>WOS:000350770500007</accession-num><urls><related-urls><url><style face="underline" font="default" size="100%"><Go to ISI>://WOS:000350770500007</style></url></related-urls></urls></record></Cite></EndNote>97. There are two main methods of measuring the rate, the first being a phylogenetic approach such as comparing sequences of a well-dated ancient sample with a modern sample. More commonly multiple modern samples are sequenced and the time separating them is estimated by calibration to an external reference, such as fossil evidence for the split between humans and chimpanzees; numerous such studies estimated the rate to be about 10-9 per site per year. The large number of generations separating the samples means that many mutations are observed, however there are a number of downsides, including the impact of ancestral polymorphism and the fact that calibration is only as good as the accuracy of the fossil date and assumed phylogenetic tree. The second approach is to use direct observation of mutations in pedigrees such as parent–offspring trios using whole-genome or exome sequencing. However, the number of de novo mutations is low, because of the small number of generations being assayed. Much debate has ensued as pedigree estimates of mutation rates are consistently about half of those using fossil calibrations (0.4–0.6 × 10-9 per nucleotide per year, using generation times between 20 and 30 years ADDIN EN.CITE <EndNote><Cite><Author>Kong</Author><Year>2012</Year><RecNum>80</RecNum><DisplayText><style face="superscript">98</style></DisplayText><record><rec-number>80</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487079810">80</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Kong, A.</author><author>Frigge, M. L.</author><author>Masson, G.</author><author>Besenbacher, S.</author><author>Sulem, P.</author><author>Magnusson, G.</author><author>Gudjonsson, S. A.</author><author>Sigurdsson, A.</author><author>Jonasdottir, A.</author><author>Wong, W. S. W.</author><author>Sigurdsson, G.</author><author>Walters, G. B.</author><author>Steinberg, S.</author><author>Helgason, H.</author><author>Thorleifsson, G.</author><author>Gudbjartsson, D. F.</author><author>Helgason, A.</author><author>Magnusson, O. T.</author><author>Thorsteinsdottir, U.</author><author>Stefansson, K.</author></authors></contributors><titles><title>Rate of de novo mutations and the importance of father's age to disease risk</title><secondary-title>Nature</secondary-title><short-title>Rate of de novo mutations and the importance of father's age to disease risk</short-title></titles><periodical><full-title>Nature</full-title></periodical><pages>471-475</pages><volume>488</volume><number>7412</number><dates><year>2012</year><pub-dates><date>Aug</date></pub-dates></dates><isbn>0028-0836</isbn><accession-num>WOS:000307761600028</accession-num><urls><related-urls><url><Go to ISI>://WOS:000307761600028</url><url>). 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ADDIN EN.CITE.DATA 99,100 which circumvents some of these caveats: heterozygous mutations found within autozygous segments must have arisen since the most recent common ancestor (MRCA), hence they provide another direct estimate of the mutation rate ADDIN EN.CITE <EndNote><Cite><Author>Broman</Author><Year>1999</Year><RecNum>25</RecNum><DisplayText><style face="superscript">3</style></DisplayText><record><rec-number>25</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487079810">25</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Broman, K. W.</author><author>Weber, J. L.</author></authors></contributors><titles><title>Long homozygous chromosomal segments in reference families from the centre d'Etude du polymorphisme humain</title><secondary-title>American Journal of Human Genetics</secondary-title><short-title>Long homozygous chromosomal segments in reference families from the centre d'Etude du polymorphisme humain</short-title></titles><periodical><full-title>American Journal of Human Genetics</full-title><abbr-1>Am. J. Hum. Genet.</abbr-1><abbr-2>Am J Hum Genet</abbr-2></periodical><pages>1493-500</pages><volume>65</volume><dates><year>1999</year></dates><label>Broman1999</label><urls><related-urls><url><style face="underline" font="default" size="100%"> face="underline" font="default" size="100%">. The number of generations to the MRCA can be inferred from the ROH length distribution and verified from pedigree information. Thus, no external calibration is required and because this approach makes use of mutations that have occurred over many generations, there is good statistical power. Care must be taken to correct for both the effects of gene conversion [G] and errors arising from the accurate calling of ROH ends. Analysis of 4,353 exome sequences from consanguineous British South AsiansPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5OYXJhc2ltaGFuPC9BdXRob3I+PFllYXI+MjAxNjwvWWVh
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ADDIN EN.CITE.DATA 41 revealed >10 Gigabases of autozygous sequences containing 932 de novo mutations, with an average of 6.6 generations separating the two copies (a little more than first cousins), providing an estimate equivalent to 0.5 × 10-9 mutations per exonic nucleotide per year, assuming 30-year generations ADDIN EN.CITE <EndNote><Cite><Author>Narasimhan</Author><Year>2015</Year><RecNum>176</RecNum><DisplayText><style face="superscript">101</style></DisplayText><record><rec-number>176</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487923889">176</key></foreign-keys><ref-type name="Conference Paper">47</ref-type><contributors><authors><author>Narasimhan, V. M.</author><author>Rahbari, R.</author><author>Scally, Aylwyn</author><author>Xue, Y.</author><author>Tyler-Smith, Chris</author><author>Durbin, R.</author></authors></contributors><titles><title>PgmNr 353: A direct estimate for the human mutation rate from autozygous sequences in thousands of parentally related pedigrees </title><secondary-title>ASHG</secondary-title></titles><num-vols>#69 The Causes and Consecuences of Evolutionary Change</num-vols><dates><year>2015</year><pub-dates><date>Oct 10, 2015</date></pub-dates></dates><pub-location>Baltimore</pub-location><urls></urls></record></Cite></EndNote>101. Approaches using any identical-by-descent haplotype sharing (including one copy) across deeper time depths provide similar estimatesPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5MaXBzb248L0F1dGhvcj48WWVhcj4yMDE1PC9ZZWFyPjxS
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ADDIN EN.CITE.DATA 102,103.Figure LegendsFigure 1 | Demographic origins of ROH. The demographic history of six diverse hypothetical populations is represented in the upper part of the plot. Representative pedigrees are indicated by dark blue lines connecting individuals (dots), loops show inbreeding, and the population size is represented by the width of the light blue areas. Thus bottlenecks are shown by a narrowing, which necessarily reduces the number of ancestral lineages that are present in the population; conversely, larger populations contain more ancestral lineages. Admixture is shown by a confluence of two hitherto separate populations, and mating between the pedigree lineages therein. The consequences of each demographic scenario are illustrated below: schematic chromosomes showing the typical distribution of runs of homozygosity (ROH) in each and at the bottom a plot of the sum total length of ROH (SROH) versus the total number of ROH (NROH) expected in each scenario. As can be seen, the burden of ROH relates to the size of the population, with smaller populations having more and longer ROH than larger populations. Admixture brings together different haplotypes and typically reduces the number of ROH to very few short ROH, whereas bottlenecks increase the number of ROH, which are typically still relatively short. Consanguinity, on the other hand, adds a small number of very long ROH for those who are the offspring of cousin marriage, thus also increasing the variance in the sum of ROH, visible as a right shift in the NROH versus SROH plot. Some populations are both bottlenecked and practice consanguineous marriage, hence having many short and some long ROH, resulting in the highest burden of ROH. Figure 2 Global census of ROH. a | Violin plot of sum length of runs of homozygosity (SROH) in 27 regional or demographic groups coloured by biogeographical continent (Americas in beige, Pacific in mint green, East Asia in pink, South Asia in mid blue, West Eurasia in orange, sub-Saharan Africa in green, and admixed in turquoise). The violin shows a coloured kernel density trace with the interquartile range as a black line and the median as a white circle. For each group, long runs of homozygosity (ROH) (>10?Mb) are plotted above and shorter ROH (2–5 Mb) are plotted below. Native Americans stand out with higher median SROH for both short and long ROH, whereas Pacific Islanders have a higher burden only of short ROH. Both West Asian and Pakistani populations have long tails in the distribution of long ROH, consistent with frequent close consanguinity. Mixed-race individuals have very few long ROH and the least short ROH. Northern Europeans are enriched for shorter ROH because the sample is mostly Finns. b | The mean SROH and number of ROH (NROH) are plotted for each of 160 populations with greater than three unrelated individuals sampled, coloured according to continent. Most populations have a complement of ROH similar to others from the same biogeographical continent, however some stand out, e.g. the Amazonian Karitiana have the highest SROH and NROH, the East African Hadza hunter-gatherers are similar to Native Greenlanders, and some North Asian groups, e.g. the Selkup are similar to Syrians and other Near Eastern populations. Populations with mean SROH>60?Mb are labelled. Published dataPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5HZW5vbWVzIFByb2plY3Q8L0F1dGhvcj48WWVhcj4yMDEy
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ADDIN EN.CITE.DATA 104-115 from the intersection of a numerous microarrays were used (147,911 single nucleotide polymorphisms (SNPs) with minor-allele frequency >0.05); individuals not clustering with their population in principal components analysis (PCA) or showing high kinship were removed before plotting; admixed Native Americans were classified using PCA and admixture analyses. Minimum ROH length 2?Mb with ≥50 SNPs. South Asians include Pakistanis, Indians, Bangladeshis, Sri Lankans and Nepalese. East Asians include Chinese, Mongolians, Japanese, Koreans, together with Southeast Asians and indigenous Siberians. Western Eurasians comprise Europeans and West Asians, which in turn include North Africans. Afr, African; Am, American; N, north; E, east; S, south; W, west; C, central; Chi, Chinese minorities; Han, Han Chinese; Jap, Japanese; Pak, Pakistani; Ind, Indian; Kh, Khoisan; Py, pygmy; Mix, mixed; Cape, Cape Coloured; Hisp, Hispanic; Lat, Latino; NatAm, Native American. Figure 3 | Genomic distributions reveal common ROH islands and random patterning of long ROH. The size and location of runs of homozygosity (ROH) over 1?Mb in length across the genome is represented by an analysis of chromosome 1 for the first 70 individuals in each of two populations from the 1000 Genomes ProjectPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5HZW5vbWVzIFByb2plY3Q8L0F1dGhvcj48WWVhcj4yMDEy
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ADDIN EN.CITE.DATA 104, together with the proportion of each population that carry ROH in each genomic location. a | Genomic distribution for Tuscans from Central Italy (Toscani in Italia; TSI). A uniform pattern of short, shared ROH in islands dominates the picture, although a few short ROH are found outside islands, due to distant pedigree loops. b | Distribution for Punjabis from Lahore, Pakistan (PJL). Again, the ROH islands stand out as concentrations of short ROH; however, in some individuals very long ROH are also present, due to the frequency of consanguineous marriage in this population. In contrast to the ROH islands, the longer ROH are distributed relatively randomly across the chromosome. c | Percentage of 92 TSI with ROH across chromosome one. There are ~10 ROH islands in Tuscans, where typically 10% to >20% of the population carries an ROH, against a background of 0.4% outside ROH islands. d | Percentage of 155 PJL with ROH. The Punjabis provide a stark contrast: the proportion of the population carrying ROH along the genome is greatly increased, averaging 2.5% outside ROH islands. The proportion of individuals in an ROH was assessed while sliding a 100?kb window across chromosome one. Windows with a red circle have a significant enrichemnt of ROH by a binomial test (P < 2 × 10-5 with Bonferroni correction for 2,500 windows).Figure 4 | Effect of genome-wide homozygosity on 16 complex traits. Significant effects are observed for two trait groups: stature-related (height and forced expiratory volume in one second (FEV1)) and cognition-related (Spearman’s g, which is a measure of generalized cognitive ability, and educational attainment)bXBlcmlhbCBDb2xsZWdlIExvbmRvbiwgTm9yZm9sayBQbGFjZSwgTG9uZG9uIFcyIDFQRywgVUsu
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ADDIN EN.CITE.DATA 20. The other, mainly cardiometabolic risk factors show no effect. BMI, body mass index; BP, blood pressure; FP fasting plasma; HbA1c, haemoglobin A1c (glycated haemoglobin); FVC, forced vital capacity; HDL, high-density lipoprotein; LDL, low-density lipoprotein.Table 1 | Studies of ROH and quantitative or disease phenotypes.PhenotypeDesignNumberPopulationResultsRefsSchizophreniaCase–Control178 cases; 144 controlsJewish ancestry, USA Individual ROH over-represented in casesPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5MZW5jejwvQXV0aG9yPjxZZWFyPjIwMDc8L1llYXI+PFJl
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ADDIN EN.CITE.DATA 31SchizophreniaCase–Control9,288 cases; 12,456 controlsMultiple countries and ancestriesAssociation with SROHPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5LZWxsZXI8L0F1dGhvcj48WWVhcj4yMDEyPC9ZZWFyPjxS
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ADDIN EN.CITE.DATA 62SchizophreniaCase–Control18,562 cases; 21,268 controlsMultiple countries and ancestriesNo association with NROH, SROH or individual ROHPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5Kb2huc29uPC9BdXRob3I+PFllYXI+MjAxNjwvWWVhcj48
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ADDIN EN.CITE.DATA 47Bipolar disorderCase–Control506 cases; 510 controlsEuropean ancestry, UKNo association with NROH, SROH or individual ROH ADDIN EN.CITE <EndNote><Cite><Author>Vine</Author><Year>2009</Year><RecNum>133</RecNum><DisplayText><style face="superscript">116</style></DisplayText><record><rec-number>133</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487079810">133</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Vine, Anna E</author><author>McQuillin, Andrew</author><author>Bass, Nicholas J</author><author>Pereira, Ana</author><author>Kandaswamy, Radhika</author><author>Robinson, Michele</author><author>Lawrence, Jacob</author><author>Anjorin, Adebayo</author><author>Sklar, Pamela</author><author>Gurling, Hugh M D</author><author>Curtis, David</author></authors></contributors><titles><title>No evidence for excess runs of homozygosity in bipolar disorder</title><secondary-title>Psychiatric Genetics</secondary-title><short-title>No evidence for excess runs of homozygosity in bipolar disorder</short-title></titles><periodical><full-title>Psychiatric Genetics</full-title><abbr-1>Psychiatr. Genet.</abbr-1><abbr-2>Psychiatr Genet</abbr-2></periodical><pages>165-70</pages><volume>19</volume><number>4</number><dates><year>2009</year></dates><isbn>1473-5873</isbn><accession-num>19451863</accession-num><label>eng</label><urls><related-urls><url><style face="underline" font="default" size="100%"> face="underline" font="default" size="100%"> by Riadh Hammami</remote-database-provider></record></Cite></EndNote>116Alzheimer’s Case–Control837 cases; 550 controlsNorth European ancestryAssociation with NROH and SROH PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5OYWxsczwvQXV0aG9yPjxZZWFyPjIwMDk8L1llYXI+PFJl
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ADDIN EN.CITE.DATA 118Autism Extreme homozygous cases5,431Multiple ancestries, USAIndividual ROH over-represented in cases ADDIN EN.CITE <EndNote><Cite><Author>Chahrour</Author><Year>2012</Year><RecNum>35</RecNum><DisplayText><style face="superscript">119</style></DisplayText><record><rec-number>35</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487079810">35</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Chahrour, M. H.</author><author>Yu, T. W.</author><author>Lim, E. T.</author><author>Ataman, B.</author><author>Coulter, M. E.</author><author>Hill, R. S.</author><author>Stevens, C. R.</author><author>Schubert, C. R.</author><author>Greenberg, M. E.</author><author>Gabriel, S. B.</author><author>Walsh, C. A.</author><author>Sequencing, Arra Autism</author></authors></contributors><titles><title>Whole-Exome Sequencing and Homozygosity Analysis Implicate Depolarization-Regulated Neuronal Genes in Autism</title><secondary-title>PLoS Genetics</secondary-title><short-title>Whole-Exome Sequencing and Homozygosity Analysis Implicate Depolarization-Regulated Neuronal Genes in Autism</short-title></titles><periodical><full-title>PLoS Genetics</full-title><abbr-1>PLoS Genet.</abbr-1><abbr-2>PLoS Genet</abbr-2></periodical><pages>236-244</pages><volume>8</volume><number>4</number><dates><year>2012</year><pub-dates><date>Apr</date></pub-dates></dates><isbn>1553-7404</isbn><accession-num>WOS:000303441800017</accession-num><urls><related-urls><url><style face="underline" font="default" size="100%"><Go to ISI>://WOS:000303441800017</style></url><url><style face="underline" font="default" size="100%"> triosMultiple ancestriesIndividual ROH over-represented in casesPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5DYXNleTwvQXV0aG9yPjxZZWFyPjIwMTI8L1llYXI+PFJl
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ADDIN EN.CITE.DATA 125Colorectal cancerCase–Control74 cases; 264 controlsJewish cases, European ancestry controls, USA Association with NROH and SROHPEVuZE5vdGU+PENpdGU+PEF1dGhvcj5CYWNvbG9kPC9BdXRob3I+PFllYXI+MjAwODwvWWVhcj48
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ADDIN EN.CITE.DATA 127Colorectal cancerCase–Control48 cases; 100 controlsSaudi ArabianNo association with NROH; individual ROH over-represented in cases ADDIN EN.CITE <EndNote><Cite><Author>Siraj</Author><Year>2012</Year><RecNum>163</RecNum><DisplayText><style face="superscript">128</style></DisplayText><record><rec-number>163</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487241806">163</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Siraj, A. K.</author><author>Khalak, H. G.</author><author>Sultana, M.</author><author>Al-Rasheed, M.</author><author>Bavi, P.</author><author>Al-Sanea, N.</author><author>Al-Dayel, F.</author><author>Uddin, S.</author><author>Alkuraya, F. S.</author><author>Al-Kuraya, K. S.</author></authors></contributors><auth-address>Human Cancer Genomic Research, Research Centre, King Faisal Specialist Hospital and Research Centre, Riyadh, Saudi Arabia.</auth-address><titles><title>Colorectal cancer risk is not associated with increased levels of homozygosity in Saudi Arabia</title><secondary-title>Genetics in Medicine</secondary-title></titles><periodical><full-title>Genetics in Medicine</full-title><abbr-1>Genet. Med.</abbr-1><abbr-2>Genet Med</abbr-2></periodical><pages>720–728</pages><dates><year>2012</year><pub-dates><date>Apr 05</date></pub-dates></dates><isbn>1530-0366 (Electronic)
1098-3600 (Linking)</isbn><accession-num>22481135</accession-num><urls><related-urls><url><style face="underline" font="default" size="100%"> acute leukaemiaCase–Control824 cases; 2,398 controlsEuropean ancestry, UKNo association with NROH, SROH or individual ROH ADDIN EN.CITE <EndNote><Cite><Author>Hosking</Author><Year>2010</Year><RecNum>65</RecNum><DisplayText><style face="superscript">129</style></DisplayText><record><rec-number>65</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487079810">65</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Hosking, F. J.</author><author>Papaemmanuil, E.</author><author>Sheridan, E.</author><author>Kinsey, S. E.</author><author>Lightfoot, T.</author><author>Roman, E.</author><author>Irving, J. A. E.</author><author>Allan, J. M.</author><author>Taylor, M.</author><author>Tomlinson, I. P.</author><author>Greaves, M.</author><author>Houlston, R. S.</author></authors></contributors><titles><title>Genome-wide homozygosity signatures and childhood acute lymphoblastic leukemia risk</title><secondary-title>Blood</secondary-title><short-title>Genome-wide homozygosity signatures and childhood acute lymphoblastic leukemia risk</short-title></titles><periodical><full-title>Blood</full-title></periodical><pages>4472-4477</pages><volume>115</volume><number>22</number><dates><year>2010</year><pub-dates><date>Jun</date></pub-dates></dates><isbn>0006-4971</isbn><accession-num>WOS:000278372600025</accession-num><urls><related-urls><url><Go to ISI>://WOS:000278372600025</url><url> & Prostate cancerCase–Control1,183 cases; 1,185 controls15 worldwide populationsNo association with NROH, SROH or individual ROH PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5FbmNpc28tTW9yYTwvQXV0aG9yPjxZZWFyPjIwMTA8L1ll
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ADDIN EN.CITE.DATA 137LongevityPopulation-based5,974DutchNo association with NROH, SROH or individual ROH ADDIN EN.CITE <EndNote><Cite><Author>Kuningas</Author><Year>2011</Year><RecNum>173</RecNum><DisplayText><style face="superscript">138</style></DisplayText><record><rec-number>173</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487749565">173</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Kuningas, M.</author><author>McQuillan, R.</author><author>Wilson, J. F.</author><author>Hofman, A.</author><author>van Duijn, C. M.</author><author>Uitterlinden, A. G.</author><author>Tiemeier, H.</author></authors></contributors><auth-address>Department of Epidemiology, Erasmus University Medical Center, Rotterdam, The Netherlands. M.Kuningas@erasmusmc.nl</auth-address><titles><title>Runs of homozygosity do not influence survival to old age</title><secondary-title>PLoS One</secondary-title></titles><periodical><full-title>PloS One</full-title><abbr-1>PLoS One</abbr-1><abbr-2>PLoS One</abbr-2></periodical><pages>e22580</pages><volume>6</volume><number>7</number><keywords><keyword>Aged</keyword><keyword>Female</keyword><keyword>*Homozygote</keyword><keyword>Humans</keyword><keyword>Longevity/*genetics</keyword><keyword>Male</keyword><keyword>Middle Aged</keyword><keyword>Polymorphism, Single Nucleotide/*genetics</keyword><keyword>Survival Analysis</keyword></keywords><dates><year>2011</year></dates><isbn>1932-6203 (Electronic)
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ADDIN EN.CITE.DATA 66Cognitive ability Population-based2,329Twins from 32 countriesNo association with NROH, SROH or individual ROH ADDIN EN.CITE <EndNote><Cite><Author>Power</Author><Year>2014</Year><RecNum>164</RecNum><DisplayText><style face="superscript">140</style></DisplayText><record><rec-number>164</rec-number><foreign-keys><key app="EN" db-id="290x02fz1se0wcez096vr0202ee2zvrrpv9s" timestamp="1487242195">164</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Power, R. A.</author><author>Nagoshi, C.</author><author>DeFries, J. C.</author><author>Plomin, R.</author><author>Wellcome Trust Case Control, Consortium</author></authors></contributors><titles><title>Genome-wide estimates of inbreeding in unrelated individuals and their association with cognitive ability</title><secondary-title>European Journal of Human Genetics</secondary-title></titles><periodical><full-title>European Journal of Human Genetics</full-title><abbr-1>Europ. J. Hum. Genet.</abbr-1><abbr-2>Europ J Hum Genet</abbr-2></periodical><pages>386-90</pages><volume>22</volume><number>3</number><keywords><keyword>Adult</keyword><keyword>Case-Control Studies</keyword><keyword>Child</keyword><keyword>*Cognition</keyword><keyword>*Consanguinity</keyword><keyword>Female</keyword><keyword>Genetic Association Studies</keyword><keyword>*Genome, Human</keyword><keyword>Homozygote</keyword><keyword>Humans</keyword><keyword>Intelligence/*genetics</keyword><keyword>Male</keyword><keyword>Polymorphism, Single Nucleotide</keyword></keywords><dates><year>2014</year><pub-dates><date>Mar</date></pub-dates></dates><isbn>1476-5438 (Electronic)
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ADDIN EN.CITE.DATA 20In this table, phenotypes are ordered by neuropsychiatric disorders, cancer, auto-immune and cardiovascular diseases, followed by quantitative traits. βFROH, the effect size estimate of FROH =1, expressed in units of intra-sex phenotypic standard deviations; NROH, number of runs of homozygosity; ROH, runs of homozygosity; SROH, sum of runs of homozygosity.Glossary termsConsanguinityMating among relatives, for example first or second cousins. Literally “of the same blood”. EndogamyMarriage within the population or community.Runs of homozygosity(ROH). Contiguous regions of the genome where an individual is homozygous across all sites. This arises if the haplotypes transmitted from the mother and father are identical, having in turn been inherited from a common ancestor at some point in the past. It is important to note this notion does not rely on a known pedigree and does not require an (arbitrary) baseline population (the first generation of ancestors or founders in a pedigree). However, ROH in practice are required to have an (arbitrary) minimum size, depending on the density of genotypes available, to distinguish identity-by-descent from chance. AutozygousHomozygosity arising at a locus due to identity-by-descent. Homozygosity-by-descent.Effective population size(Ne). The population size of an idealized population which would show the same amount of genetic drift or inbreeding, often thought of as the number of breeding individuals and usually lower than the census population size.Demographic historiesThe histories of the changes in population size, for example, populations may be large or small, of constant size, expanding or contracting, undergo bottlenecks (severe declines in population size) or founder events (establishment of populations by a limited number of ancestors), be sub-structured geographically or admix with one another.Inbreeding depressionThe reduction in evolutionary fitness of a population or individual due to the presence of increased homozygosity, arising from inbreeding. Values of traits related to fitness, such as fertility, are reduced. Genetic architectureThe make-up of the genetic basis of a trait, in particular whether there are few or many causal loci, whether the causal variants are rare or common, have small or large effect sizes, and the degree to which dominance plays a part. HaplotypeA set of alleles on a chromosome or chromosomal segment inherited from one parent — often a series of alleles at neighbouring loci which are strongly statistically associated, due to lack of recombination. Certain haplotypes may become common in the population due to natural selection or drift, until broken down over time by recombination.AdmixedGenetic admixture occurs when mating begins between two previously separate populations and individuals within the new population have a mix of haplotypes from each parental population.Inbreeding loopsAlso known as a pedigree loops, the connection in a pedigree between the maternal and paternal ancestors of an individual; the closed loops show how the same haplotypes could pass down both sides of families.Population bottleneckA severe decline in population size over a short time, or a lesser reduction over a longer time, followed by a recovery.Cosmopolitan populationsPopulations that are not isolated; typical urban populations.OverdominanceOverdominance occurs if the heterozygote trait value (phenotype) is outside the range of the trait values of the two homozygotes; also known as heterozygote advantage.Balancing selectionWhen two or more alleles are favoured by natural selection, rather than one, for example when the heterozygote is fitter than either homozygote.DominanceDominance is present at a genetic locus when the effect of one copy of an allele gives rise to a trait or phenotypic value that, rather than being half-way between the values for the two homozygotes, is nearer the trait value for a carrier of two copies of the allele. In this situation the other allele is recessive. Directional dominanceDirectional dominance occurs when the dominance effect across all causal loci in the genome has a trend in one direction, i.e. to raise or lower the trait, rather than the individual dominance effects at loci cancelling each other out. Identity-by-descent(IBD). The inheritance of an identical haplotype from both parents due to it having been passed without recombination from a common ancestor in the baseline population. Inbreeding coefficientThe probability, denoted F, of inheriting two alleles identical-by-descent at an autosomal locus, in the presence of consanguinity. F is 1/16th for first cousin offspring, 1/64th for second cousins and 1/8 for the progeny of avuncular or double first cousin matings.Genomic inbreeding coefficientThe genomic inbreeding coefficient, FROH, is the proportion of the genome that is in ROH. F and FROH have been shown to be highly correlated.Avuncular unionMarriage or mating between an uncle and neice, or aunt and nephew.ConfoundingLiterally, confusion. Statistical confounding arises when the association between a proposed explanatory variable and an outcome is distorted by the presence of a third variable associating with both. Unless all confounding can be excluded, causal inferences cannot be made from observational associations.Population stratificationA form of population genetic structure, stratification results from systematic differences in allele frequencies between subpopulations, most commonly due to differences in ancestry. If cases and controls are drawn differentially from these different subpopulations, then ancestry is a confounder and spurious associations can arise.Darwinian fitnessThe expected relative contribution of an individual or allele to the next generation of the population. It is the ability of an organism of a particular genotype to survive and leave viable offspring in its particular environment, captured in the phrase “the survival of the fittest”, although reproduction of the fittest might be more apt. PanmixiaRandom mating rather than mating structured by geography, ethnicity, socio-economic status or other factors.HeterosisAlso called hybrid vigour, the propensity when inbred lines of, for example, maize or domesticated animals are crossed that the resulting hybrids are fitter than either parent. The trait values that were reduced by inbreeding depression increase after outbreeding. Outbreeding depressionWhen the offspring of distantly related mates are less fit than the parents, for example if one homozygote has the highest fitness, outbreeding will usually increase the number of heterozygotes and thus reduce fitness.Gene conversionA mechanism of recombination where one DNA sequence is replaced by a highly homologous one, leaving the sequences identical. In mammals gene conversion tracts are usually short, between 200?bp and 1?kb.AcknowledgementsThis work was supported by the MRC Human Genetics Unit quinquennial programme grant “QTL in Health and Disease”. F.C.C. is supported by the South African National Research Foundation (NRF) and M.R. holds a South African Research Chair in Genomics and Bioinformatics of African populations hosted by the University of the Witwatersrand, funded by the Department of Science and Technology and administered by the NRF. We thank T. Gonzalez for help with figures and G. Alvarez, R. Vilas, O. Pola?ek, T. Esko, A. Wright, H. Campbell and C. Haley for helpful discussions and comments on the peting interests statementThe authors declare no competing financial interests.Online linksFURTHER INFORMATIONJim Wilson’s homepage: References ADDIN EN.REFLIST 1Cavalli-Sforza, L. L. & Bodmer, W. The Genetic of Human Populations. (W.H.Freeman & Co Ltd, San Francisco, 1978).2Bittles, A. H. & Black, M. L. Consanguinity, human evolution, and complex diseases. Proc. Natl. Acad. Sci. U. S. A. 107, 1779-1786, (2010).3Broman, K. W. & Weber, J. L. Long homozygous chromosomal segments in reference families from the centre d'Etude du polymorphisme humain. Am. J. Hum. Genet. 65, 1493-1500, (1999).This seminal study was the first to identify long ROH, showing they are common in humans. 4Jones, C. M. R. M. Atlas of world Population History. (Facts On File, New York, 1978).5Biraben, J.-N. An essay concerning mankind's demographic evolution. J. Hum. Evol. 9, 655-663, (1980).6Gunderson, R. C. 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Francisco is interested in using runs of homozygosity to decipher the demographic history of populations and the genetic basis of complex traits and diseases. Peter K. JoshiPeter Joshi a Chancellor's Fellow at the Usher Institute of Population Health Sciences and Informatics, University of Edinburgh. He started his career as a life actuary and in 2015 completed a PhD focussed on runs of homozygosity and their association with complex traits. He now works on the genomic basis of ageing and lifespan, elucidating the biomedical pathways of ageing and disease which lead to frailty and death.David W. ClarkDavid Clark in a PhD student at the University of Edinburgh, studying non-additive effects in human complex traits. He is particularly interested in using human genetic data to answer outstanding questions in evolutionary genetics, and has been lead analyst of the ROHgen consortium since 2016.Michèle RamsayMichèle Ramsay is a molecular geneticist whose research aims to shed light on the role of African population genomic variation in susceptibility to diseases, given the ethnolinguistic and environmental diversity across the continent. She is a professor in human genetics, director of the Sydney Benner Institute for Molecular Bioscience at the University of the Witwatersrand, Johannesburg, and South African Research Chair holder in Genomics and Bioinformatics of African Populations. As a member of the Human Heredity and Health in Africa Consortium (H3Africa) she leads the AWI-Gen study on the genetic and environmental contributions to obesity and cardiometabolic disease risk in six centres across four African countries.James F. WilsonJim Wilson is a professor of human genetics at the University of Edinburgh, Scotland. He leads a number of population-based studies in his native Scotland, principally in isolated populations, where he is assessing the contribution of rare variants to disease risk. Through running international consortia, his research focuses on aspects of the genetic architecture of complex traits, in particular the role of homozygosity and inbreeding depression. He also maintains his population genetic roots, recently dissecting fine-scale population genetic structure in the British Isles and Ireland. James F. Wilson’s homepage: points? The inheritance of identical haplotypes from a common ancestor creates long tracts of homozygous genotypes known as runs of homozygosity or ROH? ROH are ubiquitous across human populations, they correlate with pedigree inbreeding, larger populations have fewer, shorter ROH, whereas isolated or bottlenecked populations have more, somewhat longer ROH. Admixed groups have the fewest ROH, whereas consanguineous communities carry very long ROH. Native American populations have the highest burdens of ROH in the world.? ROH can be detected in microarray or WGS data, using either observational approaches, e.g. that implemented in PLINK or model-based approaches. Simulations show PLINK outperforms many other methods.? ROH are non-randomly distributed across the genome, being more prevalent in areas of low recombination, but are also concentrated in small regions called ROH islands.? Quantitative traits related to stature and cognition have been robustly associated with ROH burden, implying recessive variants contribute to their genetic architecture. Case-control analyses of ROH, on the other hand, appear more easily confounded by socio-economic or cultural factors.? Both mega-cohorts and special populations are now being used to investigate diverse aspects of the scope and mechanism of inbreeding depression in humans.Table of contents blurbRuns of homozygosity (ROH) are genomic regions of homozygosity where the identical maternal and paternal haplotypes are descended from a shared common ancestor, and they are well known to occur as a result of inbreeding. This article discusses strategies for detecting ROH, their underappreciated prevalence across diverse outbred populations and implications for complex traits and human disease.Subject categoriesBiological sciences / Genetics / Population genetics[URI /631/208/457]Biological sciences / Genetics / Genetic association study[URI /631/208/205]Biological sciences / Genetics / Quantitative trait[URI /631/208/480]Biological sciences / Genetics / Clinical genetics / Disease genetics[URI /631/208/2489/144]Biological sciences / Molecular biology / Chromosomes[URI /631/337/103]Biological sciences / Genetics / Genome / Genetic variation[URI /631/208/726/649] ................
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