Estimating F-statistics.

A moment estimator of, the coancestry coefficient for alleles within a population, was described by Weir & Cockerham in 1984 (100) and is still widely cited. The estimate is used by population geneticists to characterize population structure, by ecologists to estimate migration rates, by animal breeders to describe genetic variation, and by forensic scientists to quantify the strength of matching DNA profiles. This review extends the work of Weir & Cockerham by allowing different levels of coancestry for different populations, and by allowing non-zero coancestries between pairs of populations. All estimates are relative to the average value of theta between pairs of populations. Moment estimates for within- and between-population theta values are likely to have large sampling variances, although these may be reduced by combining information over loci. Variances also decrease with the numbers of alleles at a locus, and with the numbers of populations sampled. This review also extends the work of Weir & Cockerham by employing maximum likelihood methods under the assumption that allele frequencies follow the normal distribution over populations. For the case of equal theta values within populations and zero theta values between populations, the maximum likelihood estimate is the same as that given by Robertson & Hill in 1984 (70). The review concludes by relating functions of theta values to times of population divergence under a pure drift model.

[1]  Rong‐Cai Yang ESTIMATING HIERARCHICAL F‐STATISTICS , 1998, Evolution; international journal of organic evolution.

[2]  F. Viard,et al.  Crop-weed interactions in the Beta vulgaris complex at a local scale: allelic diversity and gene flow within sugar beet fields , 2002, Theoretical and Applied Genetics.

[3]  R. Fisher 014: On the "Probable Error" of a Coefficient of Correlation Deduced from a Small Sample. , 1921 .

[4]  F. Balloux,et al.  The estimation of population differentiation with microsatellite markers , 2002, Molecular ecology.

[5]  B. Weir Genetic Data Analysis II. , 1997 .

[6]  C. Saavedra,et al.  Allozyme variation in populations of scallops, Pecten jacobaeus (L.) and P. maximus (L.) (Bivalvia: Pectinidae), across the Almeria–Oran front , 2002 .

[7]  J. Long The allelic correlation structure of Gainj- and Kalam-speaking people. I. The estimation and interpretation of Wright's F-statistics. , 1986, Genetics.

[8]  N. Keyfitz The Structure of Human Populations , 1973 .

[9]  E. Krafsur Population structure of the tsetse fly Glossina pallidipes estimated by allozyme, microsatellite and mitochondrial gene diversities , 2002, Insect molecular biology.

[10]  I W Evett,et al.  Inferring ethnic origin by means of an STR profile. , 2001, Forensic science international.

[11]  Dipak K Dey,et al.  A Bayesian approach to inferring population structure from dominant markers , 2002, Molecular ecology.

[12]  M. Whitlock,et al.  Indirect measures of gene flow and migration: FST≠1/(4Nm+1) , 1999, Heredity.

[13]  T. Quinn,et al.  Reconstructing recent divergence: evaluating nonequilibrium population structure in New Zealand chinook salmon , 2002, Molecular ecology.

[14]  S. Banks,et al.  Female dispersal and relatedness structure in common wombats (Vombatus ursinus) , 2002 .

[15]  L. Jin,et al.  Ethnic-affiliation estimation by use of population-specific DNA markers. , 1997, American journal of human genetics.

[16]  L. Weigt,et al.  Genetic evidence for `leaky' cohorts in the semivoltine stonefly Peltoperla tarteri (Plecoptera: Peltoperlidae) , 2002 .

[17]  F. Balloux,et al.  Statistical properties of population differentiation estimators under stepwise mutation in a finite island model , 2002, Molecular ecology.

[18]  G. Vendramin,et al.  Origin identification of maritime pine stands in France using chloroplast simple-sequence repeats , 2002 .

[19]  K. Spitze Population structure in Daphnia obtusa: quantitative genetic and allozymic variation. , 1993, Genetics.

[20]  B. Weir,et al.  Correlations, descent measures: drift with migration and mutation. , 1987, Proceedings of the National Academy of Sciences of the United States of America.

[21]  C. Sing,et al.  A model for analysis of population structure. , 1974, Genetics.

[22]  G. Hurst,et al.  History of infection with different male-killing bacteria in the two-spot ladybird beetle Adalia bipunctata revealed through mitochondrial DNA sequence analysis. , 2002, Genetics.

[23]  Kent E. Holsinger,et al.  Analysis of Genetic Diversity in Geographically Structured Populations: A Bayesian Perspective , 2004 .

[24]  J. Anaya,et al.  Rheumatoid arthritis association in Colombian population is restricted to HLA-DRB1*04 QRRAA alleles , 2002, Genes and Immunity.

[25]  ESTIMATING THE UNBIASED ESTIMATOR θ FOR POPULATION GENETIC SURVEY DATA , 2001 .

[26]  W. Stam,et al.  The contribution of haploids, diploids and clones to fine‐scale population structure in the seaweed Cladophoropsis membranacea (Chlorophyta) , 2002, Molecular ecology.

[27]  N. Perrin,et al.  Testing demographic models of effective population size , 2001, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[28]  Kathryn Roeder,et al.  Measuring heterogeneity in forensic databases using hierarchical Bayes models , 1998 .

[29]  P. Brémond,et al.  Evolutionary aspects of the metapopulation dynamics of Biomphalaria pfeifferi, the intermediate host of Schistosoma mansoni , 2002 .

[30]  B S Weir,et al.  The effects of inbreeding on forensic calculations. , 1994, Annual review of genetics.

[31]  L. Bernatchez,et al.  Genetic structure of deep-water redfish, Sebastesmentella, populations across the North Atlantic , 2002 .

[32]  B S Weir,et al.  Estimation of the coancestry coefficient: basis for a short-term genetic distance. , 1983, Genetics.

[33]  L. Excoffier,et al.  A generic estimation of population subdivision using distances between alleles with special reference for microsatellite loci. , 1996, Genetics.

[34]  D. Couvet,et al.  Estimation of effective population size and migration rate from one- and two-locus identity measures. , 2001, Genetics.

[35]  S. Wright,et al.  Genetical Structure of Populations , 1950, Nature.

[36]  A. Gharrett,et al.  Population structure of odd‐broodline Asian pink salmon and its contrast to the even‐broodline structure , 2002 .

[37]  F. Simard,et al.  Evidence for genetic differentiation between the molecular forms M and S within the Forest chromosomal form of Anopheles gambiae in an area of sympatry , 2002, Insect molecular biology.

[38]  D. Balding,et al.  Handbook of statistical genetics , 2004 .

[39]  A. Wąs,et al.  Genetic differentiation in hatchery and wild sea trout (Salmo trutta) in the Southern Baltic at microsatellite loci , 2002 .

[40]  J. Wakeley The coalescent in an island model of population subdivision with variation among demes. , 2001, Theoretical population biology.

[41]  P. Smouse,et al.  Multivariate analysis of HLA-disease associations. , 1982, Biometrics.

[42]  C. Brenner Difficulties in the estimation of ethnic affiliation. , 1998, American journal of human genetics.

[43]  M. Whitlock,et al.  Indirect measures of gene flow and migration: FST not equal to 1/(4Nm + 1). , 1999, Heredity.

[44]  M Slatkin,et al.  A measure of population subdivision based on microsatellite allele frequencies. , 1995, Genetics.

[45]  H. Ernest,et al.  Fecal DNA Analysis and Risk Assessment of Mountain Lion Predation of Bighorn Sheep , 2002 .

[46]  J A Lambert,et al.  Genetic differentiation within and between four UK ethnic groups. , 2000, Forensic science international.

[47]  J. Curran,et al.  Conditional genotypic probabilities for microsatellite loci. , 2000, Genetics.

[48]  D J Balding,et al.  DNA profile match probability calculation: how to allow for population stratification, relatedness, database selection and single bands. , 1994, Forensic science international.

[49]  S. Engen,et al.  Inferring patterns of migration from gene frequencies under equilibrium conditions. , 1996, Genetics.

[50]  T. Reusch Microsatellites reveal high population connectivity in eelgrass (Zostera marina) in two contrasting coastal areas , 2002 .

[51]  A. K. Daníelsdóttir,et al.  Genetic heterogeneity and growth properties of different genotypes of Atlantic cod (Gadus morhua L.) at two spawning sites off south Iceland , 2002 .

[52]  L. Contu,et al.  Molecular variation of HLA class I genes in the Corsican population: approach to its origin. , 2002, European journal of immunogenetics : official journal of the British Society for Histocompatibility and Immunogenetics.

[53]  Laurent Excoffier,et al.  Analysis of Population Subdivision , 2004 .

[54]  M. Yarus Primordial genetics: phenotype of the ribocyte. , 2002, Annual review of genetics.

[55]  Eviatar Nevo,et al.  Analysis of simple sequence repeats (SSRs) in wild barley from the Fertile Crescent: associations with ecology, geography and flowering time , 2002, Plant Molecular Biology.

[56]  P. Boursot,et al.  Interpretation of variation across marker loci as evidence of selection. , 2001, Genetics.

[57]  R. Broughton,et al.  Microsatellite variation suggests substantial gene flow between king mackerel (Scomberomorus cavalla) in the western Atlantic Ocean and Gulf of Mexico , 2002 .

[58]  K. Ritland,et al.  Genetic diversity and differentiation of Kermode bear populations , 2002, Molecular ecology.

[59]  M. Iriondo,et al.  Structuring the Genetic Heterogeneity of the Basque Population: A View from Classical Polymorphisms , 2002, Human biology.

[60]  Alan Robertson,et al.  180. Note: Weighting in the Estimation of Variance Components in the Unbalanced Single Classification , 1962 .

[61]  J. Rubin,et al.  Spatial and temporal population structure of sea trout at the Island of Gotland, Sweden, delineated from mitochondrial DNA , 2002 .

[62]  Peter Donnelly,et al.  Assessing population differentiation and isolation from single‐nucleotide polymorphism data , 2002 .

[63]  I. Evett,et al.  Interpreting DNA Evidence: Statistical Genetics for Forensic Scientists , 1998 .

[64]  I W Evett,et al.  An investigation of the feasibility of inferring ethnic origin from DNA profiles. , 1992, Journal - Forensic Science Society.

[65]  G. Carvalho,et al.  The interplay between colonization history and gene flow in passively dispersing zooplankton: microsatellite analysis of rotifer resting egg banks , 2002 .

[66]  M. Van Montagu,et al.  Genetic differentiation among three neighboring Brazil-cherry (Eugenia uniflora L.) populations within the Brazilian Atlantic rain forest , 2004, Biodiversity & Conservation.

[67]  F. Rousset Inferences from Spatial Population Genetics , 2004 .

[68]  F. Rousset,et al.  AN EXACT TEST FOR POPULATION DIFFERENTIATION , 1995, Evolution; international journal of organic evolution.

[69]  K. Dodds Resampling methods in genetics and the effect of family structure in genetic data , 1986 .

[70]  C. Cockerham,et al.  ESTIMATION OF GENE FLOW FROM F‐STATISTICS , 1993, Evolution; international journal of organic evolution.

[71]  Ranajit Chakraborty,et al.  Interpreting DNA evidence , 2000 .

[72]  Bruce Budowle,et al.  Genotype Profiles for Six Population Groups at the 13 CODIS Short Tandem Repeat Core Loci and Other PCRBbased Loci , 1999 .

[73]  C. Tsigenopoulos,et al.  Biogeography and pattern of gene flow among Barbus species (Teleostei: Cyprinidae) inhabiting the Italian Peninsula and neighbouring Adriatic drainages as revealed by allozyme and mitochondrial sequence data , 2002 .

[74]  J. Pemberton,et al.  Genetic evidence for male biased dispersal in the red‐billed quelea Quelea quelea , 2002, Molecular ecology.

[75]  G Luikart,et al.  New methods employing multilocus genotypes to select or exclude populations as origins of individuals. , 1999, Genetics.

[76]  M. Monaghan,et al.  Population Genetic Structure of 3 Alpine Stream Insects: Influences of Gene Flow, Demographics, and Habitat Fragmentation , 2002, Journal of the North American Benthological Society.

[77]  P. Wolańska-Nowak Application of subpopulation theory to evaluation of DNA evidence. , 2000, Forensic science international.

[78]  B. Weir,et al.  ESTIMATING F‐STATISTICS FOR THE ANALYSIS OF POPULATION STRUCTURE , 1984, Evolution; international journal of organic evolution.

[79]  M. Endsley,et al.  Microsatellite loci in the house fly, Musca domestica L. (Diptera: Muscidae) , 2002 .

[80]  K J Dawson,et al.  A Bayesian approach to the identification of panmictic populations and the assignment of individuals. , 2001, Genetical research.

[81]  R. Bowyer,et al.  Mitochondrial phylogeography of moose (Alces alces): late pleistocene divergence and population expansion. , 2002, Molecular phylogenetics and evolution.

[82]  Creating New Paradigms of Womanhood in Modern Korean Literature: Na Hye-sok's "Kyonghui" , 2002 .

[83]  É. Fièvet,et al.  Genetic differentiation among populations of the amphidromous shrimp Atya innocous (HERBST) and obstacles to their upstreaming migration , 2002 .

[84]  J. Merilä,et al.  Comparison of genetic differentiation at marker loci and quantitative traits , 2001 .

[85]  R. Newman,et al.  FINE-SCALE POPULATION STRUCTURE IN THE WOOD FROG (RANA SYLVATICA) IN A NORTHERN WOODLAND , 2002 .

[86]  E. Geffen,et al.  Characteristics of sex‐biased dispersal and gene flow in coastal river otters: implications for natural recolonization of extirpated populations , 2002, Molecular ecology.

[87]  B. Richardson,et al.  DNA from bird‐dispersed seed and wind‐disseminated pollen provides insights into postglacial colonization and population genetic structure of whitebark pine (Pinus albicaulis) , 2002, Molecular ecology.

[88]  R. Spielman,et al.  Multivariate classification of human populations. I. Allocation of Yanomama indians to villages. , 1976, American journal of human genetics.

[89]  R. Nielsen,et al.  Distinguishing migration from isolation: a Markov chain Monte Carlo approach. , 2001, Genetics.

[90]  F. Bonhomme,et al.  Properties of bias and variance of two multiallelic estimators of F(ST). , 2000, Theoretical population biology.

[91]  D. Heath,et al.  Temporal change in genetic structure and effective population size in steelhead trout (Oncorhynchus mykiss) , 2002, Molecular ecology.

[92]  W. G. Hill,et al.  On the precision of estimation of genetic distance , 1999, Genetics Selection Evolution.

[93]  W. G. Hill,et al.  Deviations from Hardy-Weinberg proportions: sampling variances and use in estimation of inbreeding coefficients. , 1984, Genetics.

[94]  F. Rousset,et al.  Inbreeding and relatedness coefficients: what do they measure? , 2002, Heredity.

[95]  A. Blanco,et al.  Gene Flow Among Calomys Musculinus (Rodentia, Muridae) Populations in Argentina , 2004, Genetica.

[96]  C. Edward Proffitt,et al.  A Comparative Assessment of Genetic Diversity among Differently‐Aged Populations of Spartina alternif lora on Restored Versus Natural Wetlands , 2002 .

[97]  N. Frascaria-Lacoste,et al.  A generalized heterozygote deficiency assessed with microsatellites in French common ash populations , 2002, Molecular ecology.

[98]  V. Loeschcke,et al.  Genetic variation in original and colonizing Drosophila buzzatii populations analysed by microsatellite loci isolated with a new PCR screening method , 2002, Molecular ecology.

[99]  L. Bernatchez,et al.  Differential patterns of spatial divergence in microsatellite and allozyme alleles: further evidence for locus‐specific selection in the acorn barnacle, Semibalanus balanoides? , 2002, Molecular ecology.

[100]  T. F. Turner,et al.  Population structure of red drum (Sciaenops ocellatus) in the northern Gulf of Mexico, as inferred from variation in nuclear-encoded microsatellites , 2002 .