Mapping quantitative trait loci in plants: uses and caveats for evolutionary biology (original) (raw)

• Kaul, S. et al. Analysis of the genome sequence of the flowering plant Arabidopsis thaliana. Nature 408, 796– 815 (2000).
  Article CAS Google Scholar
• Theologis, A. et al. Sequence and analysis of chromosome 1 of the plant Arabidopsis thaliana. Nature 408, 816– 820 (2000).
  Article PubMed Google Scholar
• Lin, X. Y. et al. Sequence and analysis of chromosome 2 of the plant Arabidopsis thaliana. Nature 402, 761– 768 (1999).
  Article CAS PubMed Google Scholar
• Salanoubat, M. et al. Sequence and analysis of chromosome 3 of the plant Arabidopsis thaliana. Nature 408, 820– 822 (2000).
  Article CAS PubMed Google Scholar
• Mayer, K. et al. Sequence and analysis of chromosome 4 of the plant Arabidopsis thaliana. Nature 402, 769– 777 (1999).
  Article CAS PubMed Google Scholar
• Tabata, S. et al. Sequence and analysis of chromosome 5 of the plant Arabidopsis thaliana. Nature 408, 823– 826 (2000).
  Article CAS PubMed Google Scholar
• Adam, D. Now for the hard ones. Nature 408, 792– 793 (2000).
  Article CAS PubMed Google Scholar
• Bulmer, M. G. The Mathematical Theory of Quantitative Genetics (Clarendon, Oxford, 1985).
  Google Scholar
• Falconer, D. S. & Mackay, T. F. C. Introduction to Quantitative Genetics (Addison–Wesley–Longman, Harlow, 1996).
  Google Scholar
• Lynch, M. & Walsh, B. Genetics and the Analysis of Quantitative Traits (Sinauer Associates, Sunderland, Massachusetts, 1997).
  Google Scholar
• Tanksley, S. D. Mapping polygenes. Annu. Rev. Genet. 27, 205–233 (1993).
  Article CAS PubMed Google Scholar
• Liu, B.-H. Statistical Genomics: Linkage, Mapping and QTL Analysis (Boca Raton, Florida, USA, 1998).An excellent overview of QTL mapping techniques.
  Google Scholar
• Burr, B. & Burr, F. A. Recombinant inbreds for molecular mapping in maize: theoretical and practical considerations. Trends Genet. 7, 55–60 ( 1991).
  CAS PubMed Google Scholar
• Moreno-Gonzalez, J. Efficiency of generations for estimating marker-associated QTL effects by multiple regression. Genetics 135, 223– 231 (1993).
  Article CAS PubMed PubMed Central Google Scholar
• Sax, K. The association of size differences with seed-coat pattern and pigmentation in Phaseolus vulgaris. Genetics 8, 552– 560 (1923).The first mapping of a QTL by association with a Mendelian marker.
  Article CAS PubMed PubMed Central Google Scholar
• Lister, C. & Dean, C. Recombinant inbred lines for mapping RFLP and phenotypic markers in Arabidopsis thaliana. Plant J. 4, 745–750 ( 1993).
  Article CAS Google Scholar
• Cho, R. J. et al. Genome-wide mapping with biallelic markers in Arabidopsis thaliana. Nature Genet. 23, 203– 207 (1999).
  Article CAS PubMed Google Scholar
• Barton, N. H. & Turelli, M. Evolutionary quantitative genetics: how little do we know? Annu. Rev. Genet. 23, 337–370 (1989).
  Article CAS PubMed Google Scholar
• Lande, R. The response to selection on major and minor mutations affecting a metrical trait. Heredity 50, 47– 65 (1983).
  Article Google Scholar
• Orr, H. A. & Coyne, J. A. The genetics of adaptation: a reassessment . Am. Nat. 140, 725–742 (1992).The landmark paper that exhumed the argument that genes of major effect could be important in adaptation.
  Article CAS PubMed Google Scholar
• Fisher, R. A. The Genetical Theory of Natural Selection (Dover, New York, 1958).
  Google Scholar
• Alonso-Blanco, C., Blankestijn-de Vries, H., Hanhart, C. J. & Koornneef, M. Natural allelic variation at seed size loci in relation to other life history traits of Arabidopsis thaliana. Proc. Natl Acad. Sci. USA 96, 4710–4717 ( 1999).
  Article CAS PubMed PubMed Central Google Scholar
• Stanton, M. L. Seed variation in wild radish: effect of seed size on components of seedling and adult fitness. Ecology 65, 1105– 1112 (1984).
  Article Google Scholar
• Fisher, R. A. The use of multiple measurements in taxonomic problems. Ann. Eugen. 7, 179–188 ( 1936).
  Article Google Scholar
• Juenger, T., Purugganan, M. D. & Mackay, T. F. C. Quantitative trait loci for floral morphology in Arabidopsis thaliana. Genetics 156, 1379–1392 (2000).
  Article CAS PubMed PubMed Central Google Scholar
• Kelly, M. G. & Levin, D. A. Directional selection on initial flowering date in Phlox drummondii (Polemoniaceae). Am. J. Bot. 87, 382–391 ( 2000).
  Article CAS PubMed Google Scholar
• Kowalski, S. P., Lan, T. H., Feldmann, K. A. & Paterson, A. H. QTL mapping of naturally-occurring variation in flowering time of Arabidopsis thaliana. Mol. Gen. Genet. 245, 548– 555 (1994).
  Article CAS PubMed Google Scholar
• Mitchell-Olds, T. Genetic constraints on life history evolution — quantitative trait loci influencing growth and flowering in Arabidopsis thaliana. Evolution 50, 140–145 ( 1996).
  PubMed Google Scholar
• Kuittinen, H., Sillanpää, M. J. & Savolainen, O. Genetic basis of adaptation: flowering time in Arabidopsis thaliana. Theor. Appl. Genet. 95, 573–583 (1997).
  Article Google Scholar
• Byrne, M. et al. Identification and mode of action of quantitative trait loci affecting seedling height and leaf area in Eucalyptus nitens. Theor. Appl. Genet. 94, 674–681 (1997).
  Article Google Scholar
• Byrne, M., Murrell, J. C., Owen, J. V., Williams, E. R. & Moran, G. F. Mapping of quantitative trait loci influencing frost tolerance in Eucalyptus nitens. Theor. Appl. Genet. 95, 975–979 (1997).
  Article CAS Google Scholar
• Hurme, P., Silanpää, M. J., Arjas, E., Repo, T. & Savolainen, O. Genetic basis of climatic adaptation in Scots pine by Bayesian quantitative trait locus analysis. Genetics 156, 1309–1322 ( 2000).
  Article CAS PubMed PubMed Central Google Scholar
• Bradshaw, H. D. Jr & Stettler, R. Molecular genetics of growth and development in Populus. IV. Mapping QTLs with large effects on growth, form, and phenology traits in a forest tree. Genetics 139, 963– 973 (1995).
  Article CAS PubMed Google Scholar
• Frewen, B. E. et al. Quantitative trait loci and candidate gene mapping of bud set and bud flush in Populus. Genetics 154, 837–845 (2000).
  Article CAS PubMed PubMed Central Google Scholar
• Lewontin, R. C. The Genetic Basis of Evolutionary Change (Columbia University Press, New York, 1974).
  Google Scholar
• Gottlieb, L. D. Genetics and morphological evolution in plants. Am. Nat. 123, 681–709 (1984). An early, but underappreciated, paper that raised the issue that genes of major effect could be important in speciation.
  Article Google Scholar
• Coyne, J. A. & Lande, R. The genetic basis of species differences in plants. Am. Nat. 126, 141– 145 (1985).
  Article Google Scholar
• Bradshaw, H. D. Jr, Wilbert, S. M., Otto, K. G. & Schemske, D. W. Genetic mapping of floral traits associated with reproductive isolation in monkeyflowers (Mimulus). Nature 376, 762–765 ( 1995).One of the first attempts in natural populations to identify genes that are important in speciation by QTL mapping.
  Article CAS Google Scholar
• Bradshaw, H. D. Jr, Otto, K. G., Frewen, B. E., McKay, J. K. & Schemske, D. W. Quantitative trait loci affecting differences in floral morphology between two species of monkeyflower (Mimulus). Genetics 149, 367–382 (1998).
  Article CAS PubMed PubMed Central Google Scholar
• Schemske, D. W. & Bradshaw, H. D. Jr. Pollinator preference and the evolution of floral traits in monkeyflowers (Mimulus). Proc. Natl Acad. Sci. USA 96, 11910–11915 (1999).
  Article CAS PubMed PubMed Central Google Scholar
• Kelly, A. J. & Willis, J. H. Polymorphic microsatellite loci in Mimulus guttatus and related species. Mol. Ecol. 7, 769–774 (1998).
  Article CAS Google Scholar
• Cruzan, M. B. & Arnold, M. L. Ecological and genetic associations in an Iris hybrid zone. Evolution 47, 1432–1445 (1993).
  Article PubMed Google Scholar
• Arnold, M. L. Anderson's paradigm: Louisiana irises and the study of evolutionary phenomena . Mol. Ecol. 9, 1687–1698 (2000).
  Article CAS PubMed Google Scholar
• Hodges, S. A. & Arnold, M. L. Floral and ecological isolation between Aquilegia formosa and Aquilegia pubescens. Proc. Natl Acad. Sci. USA 91, 2493– 2496 (1994).
  Article CAS PubMed PubMed Central Google Scholar
• Rieseberg, L. H., Sinervo, B., Linder, C. R., Ungerer, M. C. & Arias, D. M. Role of gene interactions in hydrid speciation: evidence from ancient and experimental hybrids. Science 272, 741–745 ( 1996).
  Article CAS PubMed Google Scholar
• Kim, S.-C. & Rieseberg, L. H. Genetic architecture of species differences in annual sunflowers: implications for adaptive trait introgression . Genetics 153, 965–977 (1999).
  Article CAS PubMed PubMed Central Google Scholar
• Rieseberg, L. H., Whitton, J. & Gardner, K. Hybrid zones and the genetic architecture of a barrier to gene flow between two sunflower species. Genetics 152, 713–977 (1999).
  Article CAS PubMed PubMed Central Google Scholar
• Hodges, S. A., Burke, J. M. & Arnold, M. L. Natural formation of Iris hybrids: experimental evidence on the establishment of hybrid zones. Evolution 50, 2504–2509 (1996).
  Article PubMed Google Scholar
• Stuber, C. W. Mapping and manipulating quantitative traits in maize. Trends Genet. 11, 477–481 ( 1995).
  Article CAS PubMed Google Scholar
• Young, N. D. A cautiously optimistic vision for marker-assisted breeding. Mol. Breed. 5, 505–510 ( 1999).
  Article Google Scholar
• Grandillo, S., Ku, H. M. & Tanksley, S. D. Identifying the loci responsible for natural variation in fruit size and shape in tomato. Theor. Appl. Genet. 99, 978–987 (1999).
  Article CAS Google Scholar
• Alpert, K. B., Grandillo, S. & Tanksley, S. D. fw-2.2 — a major QTL controlling fruit weight is common to both red-fruited and green-fruited tomato species. Theor. Appl. Genet. 91, 994–1000 (1995).
  Article CAS PubMed Google Scholar
• Alpert, K. B. & Tanksley, S. D. High-resolution mapping and isolation of a yeast artificial chromosome contig containing fw2.2: a major fruit weight quantitative trait locus in tomato. Proc. Natl Acad. Sci. USA 93, 15503–15507 (1996).
  Article CAS PubMed PubMed Central Google Scholar
• Frary, A. et al. fw2.2: A quantitative trait locus key to the evolution of tomato fruit size. Science 289, 85– 88 (2000).A landmark in QTL analysis: the first molecular characterization of a locus that was originally identified entirely by QTL mapping. References 51 – 53 are the prologue to this milestone.
  Article CAS PubMed Google Scholar
• Doebley, J. & Stec, A. Genetic analysis of the morphological differences between maize and teosinte. Genetics 129 , 285–295 (1991).
  Article CAS PubMed PubMed Central Google Scholar
• Doebley, J. Mapping the genes that made maize. Trends Genet. 8, 302–307 (1992).
  Article CAS PubMed Google Scholar
• Doebley, J. & Stec, A. Inheritance of the morphological differences between maize and teosinte: comparison of results for two F2 populations . Genetics 134, 559–570 (1993).
  Article CAS PubMed PubMed Central Google Scholar
• Doebley, J., Stec, A. & Gustus, C. Teosinte branched 1 and the origin of maize: evidence for epistasis and the evolution of dominance. Genetics 141, 333–346 (1995).
  Article CAS PubMed PubMed Central Google Scholar
• White, S. & Doebley, J. Of genes and genomes and the origin of maize. Trends Genet. 14, 327– 332 (1998).
  Article CAS PubMed Google Scholar
• Dorweiler, J., Stec, A., Kermicle, J. & Doebley, J. Teosinte glume architecture 1: a genetic locus controlling a key step in maize evolution . Science 262, 233–235 (1993).
  Article CAS PubMed Google Scholar
• Doebley, J., Stec, A. & Hubbard, L. The evolution of apical dominance in maize. Nature 386, 485–488 ( 1997).
  Article CAS PubMed Google Scholar
• Wang, R. L., Stec, A., Hey, J., Lukens, L. & Doebley, J. The limits of selection during maize domestication. Nature 398, 236–239 ( 1999).References 55 – 62 chronicle a decade of work on understanding the genetics of the domestication of modern maize.
  Article CAS PubMed Google Scholar
• Paterson, A. H. et al. Convergent domestication of cereal crops by independent mutations at corresponding genetic loci. Science 269, 1714–1718 (1995). An excellent example of the power of comparative QTL mapping.
  Article CAS PubMed Google Scholar
• Ramsey, J. & Schemske, D. W. Pathways, mechanisms, and rates of polyploid formation in flowering plants. Annu. Rev. Ecol. Syst. 29, 467–501 ( 1998).
  Article Google Scholar
• Soltis, P. S. & Soltis, D. E. The role of genetic and genomic attributes in the success of polyploids. Proc. Natl Acad. Sci. USA 97, 7051–7057 ( 2000).
  Article CAS PubMed PubMed Central Google Scholar
• Hilu, K. W. Polyploidy and the evolution of domesticated plants. Am. J. Bot. 80, 1494–1499 ( 1993).
  Article Google Scholar
• Paterson, A. H. et al. Comparative genomics of plant chromosomes. Plant Cell 12, 1523–1539 ( 2000).
  Article CAS PubMed PubMed Central Google Scholar
• Jiang, C. X., Wright, R. J., El-Zik, K. M. & Paterson, A. H. Polyploid formation created unique avenues for response to selection in Gossypium (cotton). Proc. Natl Acad. Sci. USA 95 , 4419–4424 (1998).
  Article CAS PubMed PubMed Central Google Scholar
• Wright, R. J., Thaxton, P. M., El-Zik, K. M. & Paterson, A. H. D-subgenome bias of Xcm resistance genes in tetraploid Gossypium (cotton) suggests that polyploid formation has created novel avenues for evolution. Genetics 149, 1987– 1996 (1998).
  Article CAS PubMed PubMed Central Google Scholar
• Beavis, W. D. The power and deceit of QTL experiments: lessons from comparative QTL studies . Proc. Corn and Sorghum Industry Res. Conf., Am. Seed Trade Assoc., Washington DC, 255–266 ( 1994).
• Beavis, W. D. in Molecular Dissection of Complex Traits (ed. Paterson, A. H.) 145 –162 (CRC, Boca Raton, Florida, 1998 ).References 70 and 71 provided the first and most influential caveats for the use of QTL analysis in both agriculture and evolutionary biology.
  Google Scholar
• Copenhaver, G. P., Browne, W. E. & Preuss, D. Assaying genome-wide recombination and centromere functions with Arabidopsis tetrads. Proc. Natl Acad. Sci. USA 95, 247–252 (1998).
  Article CAS PubMed PubMed Central Google Scholar
• Begun, D. J. & Aquadro, C. F. Levels of naturally occurring DNA polymorphism correlate with recombination rates in D. melanogaster . Nature 356, 519–520 (1992).
  Article CAS PubMed Google Scholar
• Risch, N. J. Searching for genetic determinants in the new millennium. Nature 405, 847–856 ( 2000).
  Article CAS PubMed Google Scholar
• Weiss, K. M. & Terwilliger, J. D. How many diseases does it take to map a gene with SNPs? Nature Genet. 26, 151–157 (2000).
  Article CAS PubMed Google Scholar
• Cardon, L. R. & Bell, J. I. Association study designs for complex diseases. Nature Rev. Genet. 2, 91– 99 (2001).
  Article CAS PubMed Google Scholar
• Doerge, R. W., Zeng, Z. B. & Weir, B. S. Statistical issues in the search for genes affecting quantitative traits in experimental populations. Stat. Sci. 12, 195–219 (1997). An excellent overview of the statistical issues involved in QTL analysis.
  Article Google Scholar
• Melchinger, A. E., Utz, H. F. & Schon, C. C. Quantitative trait locus (QTL) mapping using different testers and independent population samples in maize reveals low power of QTL detection and large bias in estimates of QTL effects. Genetics 149, 383–403 ( 1998).
  Article CAS PubMed PubMed Central Google Scholar
• Otto, S. P. & Jones, C. D. Detecting the undetected: estimating the total number of loci underlying a quantitative trait. Genetics 156, 2093–2107 ( 2000).
  Article CAS PubMed PubMed Central Google Scholar
• Paterson, A. H. et al. Mendelian factors underlying quantitative traits in tomato: comparison across species, generations, and environments. Genetics 127, 181–197 ( 1991).
  Article CAS PubMed PubMed Central Google Scholar
• Jansen, R. C. Interval mapping of multiple quantitative trait loci. Genetics 135, 205–211 ( 1993).
  Article CAS PubMed PubMed Central Google Scholar
• Lander, E. & Kruglyak, L. Genetic dissection of complex traits: guidelines for interpreting and reporting linkage results. Nature Genet. 11, 241–247 ( 1995).
  Article CAS PubMed Google Scholar
• Lander, E. S. & Botstein, D. Mapping Mendelian factors underlying quantitative traits using RFLP linkage maps. Genetics 121, 185–199 (1989). In this paper, interval mapping with molecular markers to map QTL was first proposed for species in which many morphological markers were unavailable. A maximum-likelihood statistical approach for QTL mapping was also developed.
  Article CAS PubMed PubMed Central Google Scholar
• Knott, S. A. & Haley, C. S. Maximum likelihood mapping of quantitative trait loci using full-sib families. Genetics 132, 1211–1222 (1992).
  Article CAS PubMed PubMed Central Google Scholar
• Zeng, Z.-B. Precision mapping of quantitative trait loci. Genetics 136, 1457–1468 (1994).
  Article CAS PubMed PubMed Central Google Scholar
• Churchill, G. A. & Doerge, R. W. Empirical threshold values for quantitative trait mapping. Genetics 138 , 963–971 (1994).
  Article CAS PubMed PubMed Central Google Scholar
• Doerge, R. W. & Churchill, G. A. Permutation tests for multiple loci affecting a quantitative character. Genetics 142 , 285–294 (1996).
  Article CAS PubMed PubMed Central Google Scholar
• Sillanpää, M. J. & Arjas, E. Bayesian mapping of multiple quantitative trait loci from incomplete inbred line cross data . Genetics 148, 1373–1388 (1998).
  Article PubMed PubMed Central Google Scholar
• Haley, C. S., Knott, S. A. & Elsen, J.-M. Mapping quantitative trait loci in crosses between outbred lines using least squares. Genetics 136, 1195–1207 (1994).
  Article CAS PubMed PubMed Central Google Scholar
• Hoeschele, I., Uimari, P., Grignola, F. E., Zhang, Q. & Gage, K. M. Advances in statistical methods to map quantitative trait loci in outbred populations. Genetics 147, 1445–1457 ( 1997).
  Article CAS PubMed PubMed Central Google Scholar
• Sillanpää, M. J. & Arjas, E. Bayesian mapping of multiple quantitative trait loci from incomplete outbred offspring data . Genetics 151, 1605–1619 (1999).
  Article PubMed PubMed Central Google Scholar
• Pérez-Enciso, M. & Varona, L. Quantitative trait loci mapping in F2 crosses between outbred lines. Genetics 155, 391–405 ( 2000).
  Article PubMed PubMed Central Google Scholar
• George, A. W., Visscher, P. M. & Haley, C. S. Mapping quantitative trait loci in complex pedigrees: a two-step variance component approach. Genetics 156 , 2081–2092 (2000).
  Article CAS PubMed PubMed Central Google Scholar
• Hoeschele, I. in Handbook of Statistical Genetics (eds Balding, D., Bishop, M. & Cannings, C.) 599–644 (John Wiley & Sons Ltd, London, 2001).
  Google Scholar
• Liu, S.-C., Lin, Y.-R., Irvine, J. E. & Paterson, A. H. in Molecular Dissection of Complex Traits (ed. Paterson, A. H.) 95–101 (CRC, Boca Raton, Florida, 1998).
  Google Scholar
• Ming, R. et al. Detailed alignment of Saccharum and Sorghum chromosomes: comparative organization of closely related diploid and polyploid genomes . Genetics 150, 1663–1682 (1998).
  Article CAS PubMed PubMed Central Google Scholar
• Doerge, R. W. & Craig, B. A. Model selection for quantitative trait locus analysis in polyploids. Proc. Natl Acad. Sci. USA 97, 7951–7956 (2000).
  Article CAS PubMed PubMed Central Google Scholar
• Rieseberg, L. H. & Buerkle, C. A. Genetic mapping in hybrid zones. Am. Nat. (in the press).
• Provine, W. B. The Origins of Theoretical Population Genetics (Chicago Univ. Press, Illinois, 1971).A superb history of the early conflict between the 'Mendelians' and the 'biometricians', and its resolution.
  Google Scholar
• East, E. M. A Mendelian interpretation of variation that is apparently continuous. Am. Nat. 44, 65–82 ( 1910).
  Article Google Scholar
• Fisher, R. A. The correlation between relatives on the supposition of Mendelian inheritance . Trans. R. Soc. Edinb. 52, 399– 433 (1918).
  Article Google Scholar
• Lande, R. The maintenance of genetic variability by mutation in a polygenic character with linked loci. Genet. Res. 26, 221– 235 (1976).
  Article Google Scholar
• Lande, R. Quantitative genetic analysis of multivariate evolution, applied to brain:body size allometry. Evolution 33, 402– 416 (1979).
  Article PubMed Google Scholar
• Lande, R. & Arnold, S. J. The measurement of selection on correlated characters. Evolution 37, 1210 –1226 (1983).
  Article PubMed Google Scholar
• Via, S. & Lande, R. Genotype–environment interaction and the evolution of phenotypic plasticity. Evolution 39, 505–522 (1985).
  Article PubMed Google Scholar
• Turelli, M. & Barton, N. H. Dynamics of polygenic characters under selection. Theor. Popul. Biol. 38, 1–57 (1990).
  Article Google Scholar
• Barton, N. H. & Turelli, M. Natural and sexual selection on many loci. Genetics 127, 229 –255 (1991).
  Article CAS PubMed PubMed Central Google Scholar
• Turelli, M. & Barton, N. H. Genetic and statistical analyses of strong selection on polygenic traits: what, me normal? Genetics 138, 913–941 ( 1994).
  Article CAS PubMed PubMed Central Google Scholar
• Robertson, A. in Heritage from Mendel (ed. Brink, A.) 265– 280 (Wisconsin Univ. Press, Madison, Wisconsin, 1967 ).
  Google Scholar
• Haley, C. S. & Knott, S. A. A simple regression method for mapping quantitative trait loci in line crosses using flanking markers. Heredity 69, 315–324 ( 1992).
  Article CAS PubMed Google Scholar
• Thoday, J. M. Location of polygenes. Nature 191, 368– 370 (1961).
  Article Google Scholar
• Zeng, Z.-B. Theoretical basis for separation of multiple linked gene effects in mapping quantitative trait loci. Proc. Natl Acad. Sci. USA 90, 10972–10976 (1993).
  Article CAS PubMed PubMed Central Google Scholar
• Kao, C. H., Zeng, Z.-B. & Teasdale, R. D. Multiple interval mapping for quantitative trait loci. Genetics 152, 1203– 1216 (1999).
  Article CAS PubMed PubMed Central Google Scholar
• Zeng, Z.-B., Kao, C. H. & Basten, C. J. Estimating the genetic architecture of quantitative traits. Genet. Res. 74, 279– 289 (1999).
  Article CAS PubMed Google Scholar
• Basten, C. J., Weir, B. S. & Zeng, Z.-B. QTL Cartographer: A Reference Manual and Tutorial for QTL Mapping (Department of Statistics, North Carolina Univ. Press, Raleigh, North Carolina, 1995).
  Google Scholar
• Reinisch, A. J. et al. A detailed RFLP map of cotton, Gossypium hirsutum x Gossypium barbadense: chromosome organization and evolution in a disomic polyploid genome. Genetics 138, 829– 847 (1994).
  Article CAS PubMed PubMed Central Google Scholar