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F-statistics

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563: 2068:(SNPs) suggested that the genetic diversity between continental populations is even smaller and accounts for 3 to 7% A later study based on three million SNPs found that 12% of the genetic variation is found between continental populations and only 1% within them. Most of these studies have used the 2063:
It is well established that the genetic diversity among human populations is low, although the distribution of the genetic diversity was only roughly estimated. Early studies argued that 85–90% of the genetic variation is found within individuals residing in the same populations within continents
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would be the ratio of the average number of differences between pairs of chromosomes sampled within diploid individuals with the average number obtained when sampling chromosomes randomly from the population (excluding the grouping per individual). One can modify this definition and consider a
349: 412: 1084: 1261: 2552:"Similarity of the allele frequency and linkage disequilibrium pattern of single nucleotide polymorphisms in drug-related gene loci between Thai and northern East Asian populations: Implications for tagging SNP selection in Thais" 1992: 558:{\displaystyle F=1-{\frac {\operatorname {O} (f(\mathbf {Aa} ))}{\operatorname {E} (f(\mathbf {Aa} ))}}=1-{\frac {\operatorname {ObservedFrequency} (\mathbf {Aa} )}{\operatorname {ExpectedFrequency} (\mathbf {Aa} )}},\!} 189: 910: 631: 1841: 59:-statistics can also be thought of as a measure of the correlation between genes drawn at different levels of a (hierarchically) subdivided population. This correlation is influenced by several 1736: 1405: 2064:(intra-continental populations) and only an additional 10–15% is found between populations of different continents (continental populations). Later studies based on hundreds of thousands 902: 1419: 1126: 836: 808: 780: 2017: 722: 700: 2053: 1134: 1910: 1639: 1609: 1575: 1513: 1479: 1449: 1875: 1545: 674: 654: 392: 372: 3060: 3135: 1878:
grouping per sub-population instead of per individual. Population geneticists have used that idea to measure the degree of structure in a population.
344:{\displaystyle p^{2}(1-F)+pF{\text{ for }}\mathbf {AA} ;\ 2pq(1-F){\text{ for }}\mathbf {Aa} ;{\text{ and }}q^{2}(1-F)+qF{\text{ for }}\mathbf {aa} .} 3028: 1919: 3041: 2591:
Hannelius, Ulf; Salmela, Elina; Lappalainen, Tuuli; Guillot, Gilles; Lindgren, Cecilia M; Von Döbeln, Ulrika; Lahermo, Päivi; Kere, Juha (2008).
1079:{\displaystyle p={2\times \mathrm {obs} (AA)+\mathrm {obs} (Aa) \over 2\times (\mathrm {obs} (AA)+\mathrm {obs} (Aa)+\mathrm {obs} (aa))}=0.954} 2942:
Shalev, B. A.; Dvorin, A.; Herman, R.; Katz, Z.; Bornstein, S. (1991). "Long-term goose breeding for egg production and crammed liver weight".
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Bowcock, Anne M.; Kidd, Judith R.; Mountain, Joanna L.; Herbert, Joan M.; Carotenuto, Luciano; Kidd, Kenneth K.; Cavalli-Sforza, Luca (1991).
3092: 2979:"Analysis of molecular variance inferred from metric distances among DNA haplotypes: Application to human mitochondrial DNA restriction data" 2593:"Population substructure in Finland and Sweden revealed by the use of spatial coordinates and a small number of unlinked autosomal SNPs" 1527:
of two levels; one from the individual (I) to the subpopulation (S) and one from the subpopulation to the total (T). Then the total
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Wright, Sewall (1965). "The Interpretation of Population Structure by F-Statistics with Special Regard to Systems of Mating".
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are related to the amounts of heterozygosity at various levels of population structure. Together, they are called
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in a population; more specifically the expected degree of (usually) a reduction in heterozygosity when compared to
3180: 1578: 871: 3185: 2065: 2462:"The Distribution of Human Genetic Diversity: A Comparison of Mitochondrial, Autosomal, and Y-Chromosome Data" 2460:
Jorde, L.B.; Watkins, W.S.; Bamshad, M.J.; Dixon, M.E.; Ricker, C.E.; Seielstad, M.T.; Batzer, M.A. (2000).
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Mahasirimongkol, Surakameth; Chantratita, Wasun; Promso, Somying; Pasomsab, Ekawat; et al. (2006).
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Nelis, Mari; Esko, Tõnu; Mägi, Reedik; Zimprich, Fritz; et al. (2009). Fleischer, Robert C (ed.).
2685:"Genome-wide Insights into the Patterns and Determinants of Fine-Scale Population Structure in Humans" 1741:
This may be further partitioned for population substructure, and it expands according to the rules of
3281: 3155: 2121: 1524: 180: 145: 3333: 2098: 1092: 816: 788: 760: 2000: 705: 683: 3145: 1256:{\displaystyle F=1-{\frac {\mathrm {obs} (Aa)/n}{2pq}}=1-{138/1612 \over 2(0.954)(0.046)}=0.023} 3130: 50: 2246:"Genetics in geographically structured populations: Defining, estimating and interpreting FST" 2022: 3372: 3266: 3261: 3165: 1913:, causing some confusion in the scientific literature. A common definition is the following: 72: 19:
Not to be confused with the "F-test" statistic as used in general statistical inference. See
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Reich, David; Thangaraj, Kumarasamy; Patterson, Nick; Price, Alkes L.; et al. (2009).
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Lao, Oscar; Lu, Timothy T.; Nothnagel, Michael; Junge, Olaf; et al. (2008).
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Barbujani, Guido; Magagni, Arianna; Minch, Eric; Cavalli-Sforza, L. Luca (1997).
2302: 183:. In a simple two-allele system with inbreeding, the genotypic frequencies are: 3065: 394:
using heterozygotes in the above inbred population. This becomes one minus the
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Proceedings of the National Academy of Sciences of the United States of America
2131: 2108: 2103: 2093: 2069: 1987:{\displaystyle F_{ST}={\frac {\operatorname {var} (\mathbf {p} )}{p\,(1-p)}}\!} 1882: 1852: 1482: 399: 161: 100: 68: 24: 2955: 2660: 2643: 2568: 2551: 1266:
The different F-statistics look at different levels of population structure.
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from the 1960s onwards that heterozygosity in populations could be measured.
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dominants and heterozygotes to be the same, it was not until the advent of
3012: 2963: 2446: 2379: 2352: 2230: 2850:"Empirical Distributions of FST from Large-Scale Human Polymorphism Data" 395: 2816: 2549: 2928: 2361: 1516: 725: 568:
where the expected frequency at Hardy–Weinberg equilibrium is given by
114:-statistics was developed during the 1920s by the American geneticist 92: 3070: 3328: 2428: 2171: 736: 127: 88: 60: 2920: 2261: 2527: 2510: 2477: 2392: 131: 84: 46: 2590: 2683:
Biswas, Shameek; Scheinfeldt, Laura B.; Akey, Joshua M. (2009).
2644:"Correlation between Genetic and Geographic Structure in Europe" 2222: 2201:
Kulig, K (1985). "Utilization of emergency toxicology screens".
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Worked example of calculating F-statistics from genotypic data
1296:), using the above for subpopulations and averaging them; and 16:
Statistically expected level of heterozygosity in a population
728:, two alleles from a random individual of the population are 2790: 2734:"Genetic Structure of Europeans: A View from the North–East" 626:{\displaystyle \operatorname {E} (f(\mathbf {Aa} ))=2pq,\!} 2325: 1881:
Unfortunately, there is a large number of definitions for
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frequency of heterozygotes in a population divided by the
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Lewontin (1972). "The Apportionment of Human Diversity".
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Wright, S (1950). "Genetical structure of populations".
1413: 724:, respectively. It is also the probability that at any 2976: 2941: 2058: 2731: 2682: 2025: 2003: 1922: 1888: 1863: 1836:{\displaystyle 1-F=\prod _{i=0}^{i=I}(1-F_{i,i+1})\!} 1758: 1650: 1617: 1587: 1553: 1533: 1491: 1457: 1427: 1320: 1137: 1095: 913: 874: 819: 791: 763: 708: 686: 662: 642: 577: 415: 380: 360: 192: 2641: 2047: 2011: 1986: 1904: 1869: 1835: 1731:{\displaystyle 1-F_{IT}=(1-F_{IS})\,(1-F_{ST}).\!} 1730: 1633: 1603: 1569: 1539: 1507: 1473: 1443: 1399: 1255: 1120: 1078: 896: 830: 802: 774: 716: 694: 668: 648: 625: 557: 386: 366: 343: 1983: 1832: 1727: 622: 554: 3398: 1400:{\displaystyle (1-F_{IS})(1-F_{ST})=1-F_{IT},\,} 1288:is the inbreeding coefficient of an individual ( 1273:is the inbreeding coefficient of an individual ( 2332:Proceedings of the National Academy of Sciences 45:) describe the statistically expected level of 3034:Analyzing the genetic structure of populations 2977:Excoffier, L; Smouse, PE; Quattro, JM (1992). 2511:"Genetic variation, classification and 'race'" 1311:), and is calculated by solving the equation: 3086: 3061:F-statistics for Population Genetics Eco-Tool 2243: 2508: 2055:is the expected frequency of heterozygotes. 151: 2244:Holsinger, Kent E.; Weir, Bruce S. (2009). 2079:statistics or closely related statistics. 897:{\displaystyle f\left(\mathbf {Aa} \right)} 3093: 3079: 2793:"Reconstructing Indian population history" 2509:Jorde, Lynn B; Wooding, Stephen P (2004). 2203:The American Journal of Emergency Medicine 868:can be calculated, and the expectation of 3002: 2883: 2873: 2824: 2767: 2757: 2708: 2659: 2618: 2608: 2567: 2526: 2485: 2436: 2418: 2395:"An apportionment of human DNA diversity" 2369: 2351: 2269: 2029: 1964: 1698: 1396: 1117: 2292: 2019:is computed across sub-populations and 1417: 3100: 3399: 3386:Index of evolutionary biology articles 2906: 2847: 2689:The American Journal of Human Genetics 2466:The American Journal of Human Genetics 2149: 118:, who was interested in inbreeding in 3074: 3039:Wahlund effect, Wright's F-statistics 2200: 1857:A reformulation of the definition of 1414:Partition due to population structure 739:(1971) on a single population of the 374:is found by solving the equation for 3029:Shane's Simple Guide to F-Statistics 1307:) compared to the total population ( 735:For example, consider the data from 2059:Fixation index in human populations 13: 1160: 1157: 1154: 1048: 1045: 1042: 1022: 1019: 1016: 996: 993: 990: 962: 959: 956: 936: 933: 930: 578: 462: 431: 175:-statistics, and are derived from 14: 3418: 3022: 1846: 1523:Consider a population that has a 1303:is the effect of subpopulations ( 1292:) relative to the subpopulation ( 2297:. Vol. 6. pp. 381–98. 2005: 1952: 886: 883: 824: 821: 796: 793: 768: 765: 710: 688: 597: 594: 541: 538: 519: 516: 481: 478: 450: 447: 334: 331: 279: 276: 236: 233: 2970: 2935: 2900: 2841: 2784: 2725: 2676: 2635: 2584: 3186:Constructive neutral evolution 2543: 2502: 2453: 2386: 2319: 2286: 2237: 2194: 2143: 2089:Malecot's method of coancestry 2066:single-nucleotide polymorphism 2042: 2030: 1977: 1965: 1956: 1948: 1829: 1798: 1721: 1699: 1695: 1673: 1410:as shown in the next section. 1368: 1346: 1343: 1321: 1241: 1235: 1232: 1226: 1173: 1164: 1064: 1061: 1052: 1035: 1026: 1009: 1000: 986: 975: 966: 949: 940: 604: 601: 590: 584: 545: 534: 523: 512: 488: 485: 474: 468: 457: 454: 443: 437: 402:frequency of heterozygotes at 313: 301: 267: 255: 215: 203: 1: 3066:Population Structure (slides) 2137: 1121:{\displaystyle q=1-p=0.046\,} 831:{\displaystyle \mathbf {aa} } 803:{\displaystyle \mathbf {Aa} } 775:{\displaystyle \mathbf {AA} } 3136:Fisher's fundamental theorem 2875:10.1371/journal.pone.0049837 2759:10.1371/journal.pone.0005472 2303:10.1007/978-1-4684-9063-3_14 2215:10.1016/0735-6757(85)90177-9 2012:{\displaystyle \mathbf {p} } 717:{\displaystyle \mathbf {a} } 695:{\displaystyle \mathbf {A} } 7: 3161:Coefficient of relationship 2127:Coefficient of relationship 2082: 10: 3423: 2995:10.1093/genetics/131.2.479 2701:10.1016/j.ajhg.2009.04.015 1850: 404:Hardy–Weinberg equilibrium 51:Hardy–Weinberg expectation 18: 3381: 3316: 3290: 3252: 3227: 3194: 3156:Coefficient of inbreeding 3108: 2956:10.1080/00071669108417396 2661:10.1016/j.cub.2008.07.049 2569:10.1007/s10038-006-0041-1 2556:Journal of Human Genetics 2122:Coefficient of inbreeding 1277:) relative to the total ( 152:Definitions and equations 146:effective population size 3334:Evolutionary game theory 3116:Hardy–Weinberg principle 2099:Hardy-Weinberg principle 2048:{\displaystyle p\,(1-p)} 1451:can be partitioned into 1281:) population, as above; 3146:Shifting balance theory 2944:British Poultry Science 2250:Nature Reviews Genetics 3131:Linkage disequilibrium 3056:IAM based F-statistics 2610:10.1186/1471-2156-9-54 2420:10.1073/pnas.94.9.4516 2049: 2013: 1997:where the variance of 1988: 1906: 1905:{\displaystyle F_{ST}} 1871: 1837: 1797: 1732: 1635: 1634:{\displaystyle F_{ST}} 1605: 1604:{\displaystyle F_{IS}} 1571: 1570:{\displaystyle F_{IT}} 1541: 1520: 1509: 1508:{\displaystyle F_{IS}} 1475: 1474:{\displaystyle F_{ST}} 1445: 1444:{\displaystyle F_{IT}} 1401: 1257: 1122: 1080: 898: 832: 804: 776: 718: 696: 670: 650: 627: 559: 388: 368: 345: 181:inbreeding coefficient 144:can be used to define 3373:Quantitative genetics 3282:Balding–Nichols model 3267:Population bottleneck 3262:Small population size 3166:Selection coefficient 2353:10.1073/pnas.88.3.839 2050: 2014: 1989: 1907: 1872: 1838: 1771: 1733: 1636: 1606: 1572: 1542: 1510: 1476: 1446: 1421: 1402: 1258: 1123: 1081: 899: 833: 805: 777: 719: 697: 671: 651: 628: 560: 389: 369: 346: 3244:Background selection 3231:on genomic variation 3229:Effects of selection 3181:Population structure 2295:Evolutionary Biology 2023: 2001: 1920: 1886: 1861: 1756: 1648: 1615: 1585: 1551: 1531: 1525:population structure 1489: 1455: 1425: 1318: 1135: 1093: 911: 872: 817: 789: 761: 730:identical by descent 706: 684: 660: 640: 575: 413: 378: 358: 190: 122:. However, because 3407:Population genetics 3363:Population genomics 3239:Genetic hitchhiking 3126:Identity by descent 3102:Population genetics 2866:2012PLoSO...749837E 2817:10.1038/nature08365 2809:2009Natur.461..489R 2750:2009PLoSO...4.5472N 2411:1997PNAS...94.4516B 2344:1991PNAS...88..839B 2164:1950Natur.166..247W 750: 77:genetic hitchhiking 63:processes, such as 32:population genetics 3349:Landscape genetics 3044:2005-05-27 at the 2848:Elhaik, E (2012). 2045: 2009: 1984: 1902: 1867: 1833: 1743:binomial expansion 1728: 1631: 1601: 1567: 1537: 1521: 1505: 1471: 1441: 1397: 1253: 1118: 1076: 894: 866:allele frequencies 828: 800: 772: 746: 741:scarlet tiger moth 714: 692: 678:allele frequencies 666: 646: 623: 555: 384: 364: 341: 136:molecular genetics 124:complete dominance 3394: 3393: 3344:Genetic genealogy 3339:Fitness landscape 2312:978-1-4684-9065-7 1981: 1870:{\displaystyle F} 1540:{\displaystyle F} 1245: 1196: 1068: 862: 861: 813:Little spotting ( 669:{\displaystyle q} 649:{\displaystyle p} 549: 529:ExpectedFrequency 507:ObservedFrequency 492: 387:{\displaystyle F} 367:{\displaystyle F} 328: 289: 273: 245: 230: 97:natural selection 3414: 3303:J. B. S. Haldane 3095: 3088: 3081: 3072: 3071: 3017: 3016: 3006: 2974: 2968: 2967: 2939: 2933: 2932: 2904: 2898: 2897: 2887: 2877: 2845: 2839: 2838: 2828: 2803:(7263): 489–94. 2788: 2782: 2781: 2771: 2761: 2729: 2723: 2722: 2712: 2680: 2674: 2673: 2663: 2639: 2633: 2632: 2622: 2612: 2588: 2582: 2581: 2571: 2547: 2541: 2540: 2530: 2506: 2500: 2499: 2489: 2457: 2451: 2450: 2440: 2422: 2390: 2384: 2383: 2373: 2355: 2323: 2317: 2316: 2290: 2284: 2283: 2273: 2241: 2235: 2234: 2198: 2192: 2191: 2172:10.1038/166247a0 2147: 2054: 2052: 2051: 2046: 2018: 2016: 2015: 2010: 2008: 1993: 1991: 1990: 1985: 1982: 1980: 1959: 1955: 1940: 1935: 1934: 1911: 1909: 1908: 1903: 1901: 1900: 1876: 1874: 1873: 1868: 1842: 1840: 1839: 1834: 1828: 1827: 1796: 1785: 1737: 1735: 1734: 1729: 1720: 1719: 1694: 1693: 1669: 1668: 1640: 1638: 1637: 1632: 1630: 1629: 1610: 1608: 1607: 1602: 1600: 1599: 1576: 1574: 1573: 1568: 1566: 1565: 1547:, known here as 1546: 1544: 1543: 1538: 1514: 1512: 1511: 1506: 1504: 1503: 1480: 1478: 1477: 1472: 1470: 1469: 1450: 1448: 1447: 1442: 1440: 1439: 1406: 1404: 1403: 1398: 1392: 1391: 1367: 1366: 1342: 1341: 1262: 1260: 1259: 1254: 1246: 1244: 1221: 1217: 1208: 1197: 1195: 1184: 1180: 1163: 1151: 1127: 1125: 1124: 1119: 1085: 1083: 1082: 1077: 1069: 1067: 1051: 1025: 999: 978: 965: 939: 921: 903: 901: 900: 895: 893: 889: 837: 835: 834: 829: 827: 809: 807: 806: 801: 799: 781: 779: 778: 773: 771: 751: 723: 721: 720: 715: 713: 701: 699: 698: 693: 691: 675: 673: 672: 667: 655: 653: 652: 647: 632: 630: 629: 624: 600: 564: 562: 561: 556: 550: 548: 544: 526: 522: 504: 493: 491: 484: 460: 453: 429: 393: 391: 390: 385: 373: 371: 370: 365: 350: 348: 347: 342: 337: 329: 326: 300: 299: 290: 287: 282: 274: 271: 243: 239: 231: 228: 202: 201: 43:fixation indices 3422: 3421: 3417: 3416: 3415: 3413: 3412: 3411: 3397: 3396: 3395: 3390: 3377: 3312: 3286: 3248: 3232: 3230: 3223: 3190: 3121:Genetic linkage 3104: 3099: 3046:Wayback Machine 3025: 3020: 2975: 2971: 2940: 2936: 2921:10.2307/2406450 2905: 2901: 2846: 2842: 2789: 2785: 2730: 2726: 2681: 2677: 2648:Current Biology 2640: 2636: 2589: 2585: 2562:(10): 896–904. 2548: 2544: 2521:(11s): S28-33. 2515:Nature Genetics 2507: 2503: 2458: 2454: 2391: 2387: 2324: 2320: 2313: 2291: 2287: 2262:10.1038/nrg2611 2242: 2238: 2199: 2195: 2158:(4215): 247–9. 2148: 2144: 2140: 2116: 2112: 2085: 2077: 2061: 2024: 2021: 2020: 2004: 2002: 1999: 1998: 1960: 1951: 1941: 1939: 1927: 1923: 1921: 1918: 1917: 1893: 1889: 1887: 1884: 1883: 1862: 1859: 1858: 1855: 1849: 1811: 1807: 1786: 1775: 1757: 1754: 1753: 1712: 1708: 1686: 1682: 1661: 1657: 1649: 1646: 1645: 1622: 1618: 1616: 1613: 1612: 1592: 1588: 1586: 1583: 1582: 1558: 1554: 1552: 1549: 1548: 1532: 1529: 1528: 1496: 1492: 1490: 1487: 1486: 1462: 1458: 1456: 1453: 1452: 1432: 1428: 1426: 1423: 1422: 1416: 1384: 1380: 1359: 1355: 1334: 1330: 1319: 1316: 1315: 1301: 1286: 1271: 1222: 1213: 1209: 1207: 1185: 1176: 1153: 1152: 1150: 1136: 1133: 1132: 1094: 1091: 1090: 1041: 1015: 989: 979: 955: 929: 922: 920: 912: 909: 908: 904:derived : 882: 878: 873: 870: 869: 864:From this, the 820: 818: 815: 814: 792: 790: 787: 786: 764: 762: 759: 758: 757:White-spotted ( 709: 707: 704: 703: 687: 685: 682: 681: 661: 658: 657: 641: 638: 637: 593: 576: 573: 572: 537: 527: 515: 505: 503: 477: 461: 446: 430: 428: 414: 411: 410: 379: 376: 375: 359: 356: 355: 330: 327: for  325: 295: 291: 288: and  286: 275: 272: for  270: 232: 229: for  227: 197: 193: 191: 188: 187: 170: 165: 159: 154: 110:The concept of 41:(also known as 28: 17: 12: 11: 5: 3420: 3410: 3409: 3392: 3391: 3389: 3388: 3382: 3379: 3378: 3376: 3375: 3370: 3368:Phylogeography 3365: 3360: 3358:Microevolution 3355: 3346: 3341: 3336: 3331: 3326: 3320: 3318: 3317:Related topics 3314: 3313: 3311: 3310: 3305: 3300: 3294: 3292: 3288: 3287: 3285: 3284: 3279: 3274: 3272:Founder effect 3269: 3264: 3258: 3256: 3250: 3249: 3247: 3246: 3241: 3235: 3233: 3228: 3225: 3224: 3222: 3221: 3216: 3211: 3206: 3200: 3198: 3192: 3191: 3189: 3188: 3183: 3178: 3173: 3168: 3163: 3158: 3153: 3151:Price equation 3148: 3143: 3141:Neutral theory 3138: 3133: 3128: 3123: 3118: 3112: 3110: 3106: 3105: 3098: 3097: 3090: 3083: 3075: 3069: 3068: 3063: 3058: 3053: 3048: 3036: 3031: 3024: 3023:External links 3021: 3019: 3018: 2969: 2934: 2915:(3): 395–420. 2899: 2860:(11): e49837. 2840: 2783: 2724: 2695:(5): 641–650. 2675: 2654:(16): 1241–8. 2634: 2583: 2542: 2528:10.1038/ng1435 2501: 2478:10.1086/302825 2452: 2385: 2318: 2311: 2285: 2236: 2193: 2141: 2139: 2136: 2135: 2134: 2132:Fixation index 2129: 2124: 2119: 2114: 2110: 2106: 2104:Wahlund effect 2101: 2096: 2094:Heterozygosity 2091: 2084: 2081: 2073: 2060: 2057: 2044: 2041: 2038: 2035: 2032: 2028: 2007: 1995: 1994: 1979: 1976: 1973: 1970: 1967: 1963: 1958: 1954: 1950: 1947: 1944: 1938: 1933: 1930: 1926: 1899: 1896: 1892: 1866: 1853:Fixation index 1851:Main article: 1848: 1847:Fixation index 1845: 1844: 1843: 1831: 1826: 1823: 1820: 1817: 1814: 1810: 1806: 1803: 1800: 1795: 1792: 1789: 1784: 1781: 1778: 1774: 1770: 1767: 1764: 1761: 1745:, so that for 1739: 1738: 1726: 1723: 1718: 1715: 1711: 1707: 1704: 1701: 1697: 1692: 1689: 1685: 1681: 1678: 1675: 1672: 1667: 1664: 1660: 1656: 1653: 1628: 1625: 1621: 1598: 1595: 1591: 1564: 1561: 1557: 1536: 1502: 1499: 1495: 1483:Wahlund effect 1468: 1465: 1461: 1438: 1435: 1431: 1415: 1412: 1408: 1407: 1395: 1390: 1387: 1383: 1379: 1376: 1373: 1370: 1365: 1362: 1358: 1354: 1351: 1348: 1345: 1340: 1337: 1333: 1329: 1326: 1323: 1299: 1284: 1269: 1264: 1263: 1252: 1249: 1243: 1240: 1237: 1234: 1231: 1228: 1225: 1220: 1216: 1212: 1206: 1203: 1200: 1194: 1191: 1188: 1183: 1179: 1175: 1172: 1169: 1166: 1162: 1159: 1156: 1149: 1146: 1143: 1140: 1129: 1128: 1116: 1113: 1110: 1107: 1104: 1101: 1098: 1087: 1086: 1075: 1072: 1066: 1063: 1060: 1057: 1054: 1050: 1047: 1044: 1040: 1037: 1034: 1031: 1028: 1024: 1021: 1018: 1014: 1011: 1008: 1005: 1002: 998: 995: 992: 988: 985: 982: 977: 974: 971: 968: 964: 961: 958: 954: 951: 948: 945: 942: 938: 935: 932: 928: 925: 919: 916: 892: 888: 885: 881: 877: 860: 859: 856: 853: 850: 847: 843: 842: 839: 826: 823: 811: 798: 795: 785:Intermediate ( 783: 770: 767: 755: 712: 690: 665: 645: 634: 633: 621: 618: 615: 612: 609: 606: 603: 599: 596: 592: 589: 586: 583: 580: 566: 565: 553: 547: 543: 540: 536: 533: 530: 525: 521: 518: 514: 511: 508: 502: 499: 496: 490: 487: 483: 480: 476: 473: 470: 467: 464: 459: 456: 452: 449: 445: 442: 439: 436: 433: 427: 424: 421: 418: 383: 363: 354:The value for 352: 351: 340: 336: 333: 324: 321: 318: 315: 312: 309: 306: 303: 298: 294: 285: 281: 278: 269: 266: 263: 260: 257: 254: 251: 248: 242: 238: 235: 226: 223: 220: 217: 214: 211: 208: 205: 200: 196: 168: 163: 157: 156:The measures F 153: 150: 101:Wahlund effect 69:founder effect 47:heterozygosity 25:F-distribution 15: 9: 6: 4: 3: 2: 3419: 3408: 3405: 3404: 3402: 3387: 3384: 3383: 3380: 3374: 3371: 3369: 3366: 3364: 3361: 3359: 3356: 3354: 3350: 3347: 3345: 3342: 3340: 3337: 3335: 3332: 3330: 3327: 3325: 3322: 3321: 3319: 3315: 3309: 3308:Sewall Wright 3306: 3304: 3301: 3299: 3296: 3295: 3293: 3289: 3283: 3280: 3278: 3275: 3273: 3270: 3268: 3265: 3263: 3260: 3259: 3257: 3255: 3254:Genetic drift 3251: 3245: 3242: 3240: 3237: 3236: 3234: 3226: 3220: 3217: 3215: 3212: 3210: 3207: 3205: 3202: 3201: 3199: 3197: 3193: 3187: 3184: 3182: 3179: 3177: 3174: 3172: 3169: 3167: 3164: 3162: 3159: 3157: 3154: 3152: 3149: 3147: 3144: 3142: 3139: 3137: 3134: 3132: 3129: 3127: 3124: 3122: 3119: 3117: 3114: 3113: 3111: 3107: 3103: 3096: 3091: 3089: 3084: 3082: 3077: 3076: 3073: 3067: 3064: 3062: 3059: 3057: 3054: 3052: 3049: 3047: 3043: 3040: 3037: 3035: 3032: 3030: 3027: 3026: 3014: 3010: 3005: 3000: 2996: 2992: 2989:(2): 479–91. 2988: 2984: 2980: 2973: 2965: 2961: 2957: 2953: 2949: 2945: 2938: 2930: 2926: 2922: 2918: 2914: 2910: 2903: 2895: 2891: 2886: 2881: 2876: 2871: 2867: 2863: 2859: 2855: 2851: 2844: 2836: 2832: 2827: 2822: 2818: 2814: 2810: 2806: 2802: 2798: 2794: 2787: 2779: 2775: 2770: 2765: 2760: 2755: 2751: 2747: 2743: 2739: 2735: 2728: 2720: 2716: 2711: 2706: 2702: 2698: 2694: 2690: 2686: 2679: 2671: 2667: 2662: 2657: 2653: 2649: 2645: 2638: 2630: 2626: 2621: 2616: 2611: 2606: 2602: 2598: 2594: 2587: 2579: 2575: 2570: 2565: 2561: 2557: 2553: 2546: 2538: 2534: 2529: 2524: 2520: 2516: 2512: 2505: 2497: 2493: 2488: 2483: 2479: 2475: 2472:(3): 979–88. 2471: 2467: 2463: 2456: 2448: 2444: 2439: 2434: 2430: 2426: 2421: 2416: 2412: 2408: 2405:(9): 4516–9. 2404: 2400: 2396: 2389: 2381: 2377: 2372: 2367: 2363: 2359: 2354: 2349: 2345: 2341: 2338:(3): 839–43. 2337: 2333: 2329: 2322: 2314: 2308: 2304: 2300: 2296: 2289: 2281: 2277: 2272: 2267: 2263: 2259: 2256:(9): 639–50. 2255: 2251: 2247: 2240: 2232: 2228: 2224: 2220: 2216: 2212: 2208: 2204: 2197: 2189: 2185: 2181: 2177: 2173: 2169: 2165: 2161: 2157: 2153: 2146: 2142: 2133: 2130: 2128: 2125: 2123: 2120: 2118: 2107: 2105: 2102: 2100: 2097: 2095: 2092: 2090: 2087: 2086: 2080: 2078: 2076: 2072: 2067: 2056: 2039: 2036: 2033: 2026: 1974: 1971: 1968: 1961: 1945: 1942: 1936: 1931: 1928: 1924: 1916: 1915: 1914: 1912: 1897: 1894: 1890: 1879: 1864: 1854: 1824: 1821: 1818: 1815: 1812: 1808: 1804: 1801: 1793: 1790: 1787: 1782: 1779: 1776: 1772: 1768: 1765: 1762: 1759: 1752: 1751: 1750: 1748: 1744: 1724: 1716: 1713: 1709: 1705: 1702: 1690: 1687: 1683: 1679: 1676: 1670: 1665: 1662: 1658: 1654: 1651: 1644: 1643: 1642: 1626: 1623: 1619: 1596: 1593: 1589: 1580: 1562: 1559: 1555: 1534: 1526: 1518: 1500: 1497: 1493: 1484: 1466: 1463: 1459: 1436: 1433: 1429: 1420: 1411: 1393: 1388: 1385: 1381: 1377: 1374: 1371: 1363: 1360: 1356: 1352: 1349: 1338: 1335: 1331: 1327: 1324: 1314: 1313: 1312: 1310: 1306: 1302: 1295: 1291: 1287: 1280: 1276: 1272: 1250: 1247: 1238: 1229: 1223: 1218: 1214: 1210: 1204: 1201: 1198: 1192: 1189: 1186: 1181: 1177: 1170: 1167: 1147: 1144: 1141: 1138: 1131: 1130: 1114: 1111: 1108: 1105: 1102: 1099: 1096: 1089: 1088: 1073: 1070: 1058: 1055: 1038: 1032: 1029: 1012: 1006: 1003: 983: 980: 972: 969: 952: 946: 943: 926: 923: 917: 914: 907: 906: 905: 890: 879: 875: 867: 857: 854: 851: 848: 845: 844: 840: 812: 784: 756: 753: 752: 749: 744: 742: 738: 733: 731: 727: 679: 663: 643: 619: 616: 613: 610: 607: 587: 581: 571: 570: 569: 551: 531: 528: 509: 506: 500: 497: 494: 471: 465: 440: 434: 425: 422: 419: 416: 409: 408: 407: 405: 401: 397: 381: 361: 338: 322: 319: 316: 310: 307: 304: 296: 292: 283: 264: 261: 258: 252: 249: 246: 240: 224: 221: 218: 212: 209: 206: 198: 194: 186: 185: 184: 182: 178: 174: 166: 149: 147: 143: 139: 137: 133: 129: 125: 121: 117: 116:Sewall Wright 113: 108: 106: 105:genetic drift 102: 98: 94: 90: 86: 82: 81:meiotic drive 78: 74: 70: 66: 65:genetic drift 62: 58: 54: 52: 48: 44: 40: 38: 33: 26: 22: 3324:Biogeography 3298:R. A. Fisher 3176:Heritability 3109:Key concepts 2986: 2982: 2972: 2950:(4): 703–9. 2947: 2943: 2937: 2912: 2908: 2902: 2857: 2853: 2843: 2800: 2796: 2786: 2744:(5): e5472. 2741: 2737: 2727: 2692: 2688: 2678: 2651: 2647: 2637: 2600: 2597:BMC Genetics 2596: 2586: 2559: 2555: 2545: 2518: 2514: 2504: 2469: 2465: 2455: 2402: 2398: 2388: 2335: 2331: 2321: 2294: 2288: 2253: 2249: 2239: 2209:(6): 573–4. 2206: 2202: 2196: 2155: 2151: 2145: 2074: 2070: 2062: 1996: 1880: 1856: 1749:partitions: 1746: 1740: 1522: 1409: 1308: 1304: 1297: 1293: 1289: 1282: 1278: 1274: 1267: 1265: 863: 747: 734: 635: 567: 353: 176: 172: 155: 141: 140: 111: 109: 61:evolutionary 56: 55: 42: 36: 35: 29: 3277:Coalescence 1579:partitioned 1481:due to the 126:causes the 39:-statistics 3219:Ecological 3209:Artificial 2138:References 1517:inbreeding 132:homozygote 128:phenotypes 93:inbreeding 73:bottleneck 3329:Evolution 3196:Selection 2909:Evolution 2037:− 1972:− 1946:⁡ 1805:− 1773:∏ 1763:− 1706:− 1680:− 1655:− 1577:, can be 1378:− 1353:− 1328:− 1205:− 1148:− 1106:− 984:× 927:× 754:Genotype 748:Table 1: 737:E.B. Ford 582:⁡ 532:⁡ 510:⁡ 501:− 466:⁡ 435:⁡ 426:− 308:− 262:− 210:− 99:, or the 89:gene flow 3401:Category 3353:genomics 3291:Founders 3042:Archived 2983:Genetics 2894:23185452 2854:PLOS ONE 2835:19779445 2778:19424496 2738:PLOS ONE 2719:19442770 2670:18691889 2629:18713460 2578:16957813 2537:15508000 2496:10712212 2280:19687804 2223:67025533 2188:36311175 2180:15439261 2117:analyses 2083:See also 676:are the 400:expected 396:observed 85:mutation 3204:Natural 3171:Fitness 3013:1644282 3004:1205020 2964:1933444 2929:2406450 2885:3504095 2862:Bibcode 2826:2842210 2805:Bibcode 2769:2675054 2746:Bibcode 2710:2681007 2620:2527025 2487:1288178 2447:9114021 2407:Bibcode 2380:1992475 2362:2356081 2340:Bibcode 2271:4687486 2231:4063030 2160:Bibcode 1515:due to 846:Number 167:, and F 3214:Sexual 3011:  3001:  2962:  2927:  2892:  2882:  2833:  2823:  2797:Nature 2776:  2766:  2717:  2707:  2668:  2627:  2617:  2603:: 54. 2576:  2535:  2494:  2484:  2445:  2435:  2427:  2378:  2368:  2360:  2309:  2278:  2268:  2229:  2221:  2186:  2178:  2152:Nature 841:Total 636:where 244:  179:, the 120:cattle 21:F-test 2925:JSTOR 2438:20754 2429:42042 2425:JSTOR 2371:50909 2358:JSTOR 2184:S2CID 1581:into 1251:0.023 1239:0.046 1230:0.954 1115:0.046 1074:0.954 858:1612 849:1469 726:locus 3351:and 3009:PMID 2960:PMID 2890:PMID 2831:PMID 2774:PMID 2715:PMID 2666:PMID 2625:PMID 2574:PMID 2533:PMID 2492:PMID 2443:PMID 2376:PMID 2307:ISBN 2276:PMID 2227:PMID 2219:LCCN 2176:PMID 1611:and 1485:and 1219:1612 852:138 702:and 656:and 23:and 2999:PMC 2991:doi 2987:131 2952:doi 2917:doi 2880:PMC 2870:doi 2821:PMC 2813:doi 2801:461 2764:PMC 2754:doi 2705:PMC 2697:doi 2656:doi 2615:PMC 2605:doi 2564:doi 2523:doi 2482:PMC 2474:doi 2433:PMC 2415:doi 2366:PMC 2348:doi 2299:doi 2266:PMC 2258:doi 2211:doi 2168:doi 2156:166 1943:var 1211:138 680:of 130:of 30:In 3403:: 3007:. 2997:. 2985:. 2981:. 2958:. 2948:32 2946:. 2923:. 2913:19 2911:. 2888:. 2878:. 2868:. 2856:. 2852:. 2829:. 2819:. 2811:. 2799:. 2795:. 2772:. 2762:. 2752:. 2740:. 2736:. 2713:. 2703:. 2693:84 2691:. 2687:. 2664:. 2652:18 2650:. 2646:. 2623:. 2613:. 2599:. 2595:. 2572:. 2560:51 2558:. 2554:. 2531:. 2519:36 2517:. 2513:. 2490:. 2480:. 2470:66 2468:. 2464:. 2441:. 2431:. 2423:. 2413:. 2403:94 2401:. 2397:. 2374:. 2364:. 2356:. 2346:. 2336:88 2334:. 2330:. 2305:. 2274:. 2264:. 2254:10 2252:. 2248:. 2225:. 2217:. 2205:. 2182:. 2174:. 2166:. 2154:. 2115:ST 2113:-F 2111:ST 2075:ST 1641:: 1300:ST 1285:IS 1270:IT 855:5 838:) 810:) 782:) 743:: 732:. 406:: 169:IT 164:ST 160:, 158:IS 148:. 107:. 95:, 91:, 87:, 83:, 79:, 75:, 71:, 67:, 53:. 34:, 3094:e 3087:t 3080:v 3015:. 2993:: 2966:. 2954:: 2931:. 2919:: 2896:. 2872:: 2864:: 2858:7 2837:. 2815:: 2807:: 2780:. 2756:: 2748:: 2742:4 2721:. 2699:: 2672:. 2658:: 2631:. 2607:: 2601:9 2580:. 2566:: 2539:. 2525:: 2498:. 2476:: 2449:. 2417:: 2409:: 2382:. 2350:: 2342:: 2315:. 2301:: 2282:. 2260:: 2233:. 2213:: 2207:3 2190:. 2170:: 2162:: 2109:Q 2071:F 2043:) 2040:p 2034:1 2031:( 2027:p 2006:p 1978:) 1975:p 1969:1 1966:( 1962:p 1957:) 1953:p 1949:( 1937:= 1932:T 1929:S 1925:F 1898:T 1895:S 1891:F 1865:F 1830:) 1825:1 1822:+ 1819:i 1816:, 1813:i 1809:F 1802:1 1799:( 1794:I 1791:= 1788:i 1783:0 1780:= 1777:i 1769:= 1766:F 1760:1 1747:I 1725:. 1722:) 1717:T 1714:S 1710:F 1703:1 1700:( 1696:) 1691:S 1688:I 1684:F 1677:1 1674:( 1671:= 1666:T 1663:I 1659:F 1652:1 1627:T 1624:S 1620:F 1597:S 1594:I 1590:F 1563:T 1560:I 1556:F 1535:F 1519:. 1501:S 1498:I 1494:F 1467:T 1464:S 1460:F 1437:T 1434:I 1430:F 1394:, 1389:T 1386:I 1382:F 1375:1 1372:= 1369:) 1364:T 1361:S 1357:F 1350:1 1347:( 1344:) 1339:S 1336:I 1332:F 1325:1 1322:( 1309:T 1305:S 1298:F 1294:S 1290:I 1283:F 1279:T 1275:I 1268:F 1248:= 1242:) 1236:( 1233:) 1227:( 1224:2 1215:/ 1202:1 1199:= 1193:q 1190:p 1187:2 1182:n 1178:/ 1174:) 1171:a 1168:A 1165:( 1161:s 1158:b 1155:o 1145:1 1142:= 1139:F 1112:= 1109:p 1103:1 1100:= 1097:q 1071:= 1065:) 1062:) 1059:a 1056:a 1053:( 1049:s 1046:b 1043:o 1039:+ 1036:) 1033:a 1030:A 1027:( 1023:s 1020:b 1017:o 1013:+ 1010:) 1007:A 1004:A 1001:( 997:s 994:b 991:o 987:( 981:2 976:) 973:a 970:A 967:( 963:s 960:b 957:o 953:+ 950:) 947:A 944:A 941:( 937:s 934:b 931:o 924:2 918:= 915:p 891:) 887:a 884:A 880:( 876:f 825:a 822:a 797:a 794:A 769:A 766:A 711:a 689:A 664:q 644:p 620:, 617:q 614:p 611:2 608:= 605:) 602:) 598:a 595:A 591:( 588:f 585:( 579:E 552:, 546:) 542:a 539:A 535:( 524:) 520:a 517:A 513:( 498:1 495:= 489:) 486:) 482:a 479:A 475:( 472:f 469:( 463:E 458:) 455:) 451:a 448:A 444:( 441:f 438:( 432:O 423:1 420:= 417:F 382:F 362:F 339:. 335:a 332:a 323:F 320:q 317:+ 314:) 311:F 305:1 302:( 297:2 293:q 284:; 280:a 277:A 268:) 265:F 259:1 256:( 253:q 250:p 247:2 241:; 237:A 234:A 225:F 222:p 219:+ 216:) 213:F 207:1 204:( 199:2 195:p 177:F 173:F 162:F 142:F 112:F 57:F 37:F 27:.

Index

F-test
F-distribution
population genetics
heterozygosity
Hardy–Weinberg expectation
evolutionary
genetic drift
founder effect
bottleneck
genetic hitchhiking
meiotic drive
mutation
gene flow
inbreeding
natural selection
Wahlund effect
genetic drift
Sewall Wright
cattle
complete dominance
phenotypes
homozygote
molecular genetics
effective population size
FST
inbreeding coefficient
observed
expected
Hardy–Weinberg equilibrium
allele frequencies

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