Calvin O. Qualset

Genome comparisons reveal a dominant mechanism of chromosome number reduction in grasses and accelerated genome evolution in Triticeae

Single-nucleotide polymorphism was used in the construction of an expressed sequence tag map of Aegilops tauschii, the diploid source of the wheat D genome. Comparisons of the map with the rice and sorghum genome sequences revealed 50 inversions and translocations; 2, 8, and 40 were assigned respectively to the rice, sorghum, and Ae. tauschii lineages, showing greatly accelerated genome evolution in the large Triticeae genomes. The reduction of the basic chromosome number from 12 to 7 in the Triticeae has taken place by a process during which an entire chromosome is inserted by its telomeres into a break in the centromeric region of another chromosome. The original centromere–telomere polarity of the chromosome arms is maintained in the new chromosome. An intrachromosomal telomere–telomere fusion resulting in a pericentric translocation of a chromosome segment or an entire arm accompanied or preceded the chromosome insertion in some instances. Insertional dysploidy has been recorded in three grass subfamilies and appears to be the dominant mechanism of basic chromosome number reduction in grasses. A total of 64% and 66% of Ae. tauschii genes were syntenic with sorghum and rice genes, respectively. Synteny was reduced in the vicinity of the termini of modern Ae. tauschii chromosomes but not in the vicinity of the ancient termini embedded in the Ae. tauschii chromosomes, suggesting that the dependence of synteny erosion on gene location along the centromere–telomere axis either evolved recently in the Triticeae phylogenetic lineage or its evolution was recently accelerated.

Synteny perturbations between wheat homoeologous chromosomes caused by locus duplications and deletions correlate with recombination rates

Loci detected by Southern blot hybridization of 3,977 expressed sequence tag unigenes were mapped into 159 chromosome bins delineated by breakpoints of a series of overlapping deletions. These data were used to assess synteny levels along homoeologous chromosomes of the wheat A, B, and D genomes, in relation to both bin position on the centromere-telomere axis and the gradient of recombination rates along chromosome arms. Synteny level decreased with the distance of a chromosome region from the centromere. It also decreased with an increase in recombination rates along the average chromosome arm. There were twice as many unique loci in the B genome than in the A and D genomes, and synteny levels between the B genome chromosomes and the A and D genome homoeologues were lower than those between the A and D genome homoeologues. These differences among the wheat genomes were attributed to differences in the mating systems of wheat diploid ancestors. Synteny perturbations were characterized in 31 paralogous sets of loci with perturbed synteny. Both insertions and deletions of loci were detected and both preferentially occurred in high recombination regions of chromosomes.

Nucleotide diversity maps reveal variation in diversity among wheat genomes and chromosomes.

BackgroundA genome-wide assessment of nucleotide diversity in a polyploid species must minimize the inclusion of homoeologous sequences into diversity estimates and reliably allocate individual haplotypes into their respective genomes. The same requirements complicate the development and deployment of single nucleotide polymorphism (SNP) markers in polyploid species. We report here a strategy that satisfies these requirements and deploy it in the sequencing of genes in cultivated hexaploid wheat (Triticum aestivum, genomes AABBDD) and wild tetraploid wheat (Triticum turgidum ssp. dicoccoides, genomes AABB) from the putative site of wheat domestication in Turkey. Data are used to assess the distribution of diversity among and within wheat genomes and to develop a panel of SNP markers for polyploid wheat.ResultsNucleotide diversity was estimated in 2114 wheat genes and was similar between the A and B genomes and reduced in the D genome. Within a genome, diversity was diminished on some chromosomes. Low diversity was always accompanied by an excess of rare alleles. A total of 5,471 SNPs was discovered in 1791 wheat genes. Totals of 1,271, 1,218, and 2,203 SNPs were discovered in 488, 463, and 641 genes of wheat putative diploid ancestors, T. urartu, Aegilops speltoides, and Ae. tauschii, respectively. A public database containing genome-specific primers, SNPs, and other information was constructed. A total of 987 genes with nucleotide diversity estimated in one or more of the wheat genomes was placed on an Ae. tauschii genetic map, and the map was superimposed on wheat deletion-bin maps. The agreement between the maps was assessed.ConclusionsIn a young polyploid, exemplified by T. aestivum, ancestral species are the primary source of genetic diversity. Low effective recombination due to self-pollination and a genetic mechanism precluding homoeologous chromosome pairing during polyploid meiosis can lead to the loss of diversity from large chromosomal regions. The net effect of these factors in T. aestivum is large variation in diversity among genomes and chromosomes, which impacts the development of SNP markers and their practical utility. Accumulation of new mutations in older polyploid species, such as wild emmer, results in increased diversity and its more uniform distribution across the genome.

Genetic aspects of wheat gliadin proteins

Inheritance of gliadin components unique to three different varieties of common wheat (Triticum aestivum L.) was studied in F1 and F2 seeds of intervarietal crosses using protein patterns obtained by polyacrylamide gel electrophoresis in aluminum lactate buffer (pH 3.2). The patterns of F1 seeds of the crosses Cheyenne × Justin and INIA 66R × Justin evidenced all the bands present in the patterns of the parents; band intensities reflected gene dosage levels dependent on whether the contributing parent was maternal or paternal in accordance with the triploid nature of endosperm tissue. Most of the gliadin components examined segregated in accordance with control by a single dominant gene, but in two instances single bands in the one-dimensional electrophoretic patterns segregated in the F2 as expected if controlled by two genes. A method of two-dimensional electrophoresis was developed that resolved these apparently single bands into two components each, which could segregate independently. Linkage analysis provided evidence of codominant alleles and closely linked genes coding for gliadin protein components in both coupling and repulsion situations. The gliadin protein components seem to be coded for by clusters of genes located on chromosomes of homoeologous groups 1 and 6 in hexaploid wheats.

Quantitative trait locus analysis of wheat quality traits

Milling and baking quality traits in wheat (Triticum aestivum L.) were studied by QTL analysis in the ITMI population, a set of 114 recombinant inbred lines (RILs) generated from a synthetic-hexaploid (W7985) × bread-wheat (Opata 85) cross. Grain from RILs grown in U.S., French, and Mexican wheat-growing regions was assayed for kernel-texture traits, protein concentration and quality, and dough strength and mixing traits. Only kernel-texture traits showed similar genetic control in all environments, with Opata ha alleles at the hardness locus Ha on chromosome arm 5DS increasing grain hardness, alkaline water retention capacity, and flour yield. Dough strength was most strongly influenced by Opata alleles at 5DS loci near or identical to Ha. Grain protein concentration was associated not with high-molecular-weight glutenin loci but most consistently with the Gli-D2 gliadin locus on chromosome arm 6DS. In Mexican-grown material, a 2DS locus near photoperiod-sensitivity gene Ppd1 accounted for 25% of variation in protein, with the ppd1-coupled allele associated with higher (1.1%) protein concentration. Mixogram traits showed most influence from chromosomal regions containing gliadin or low-molecular-weight glutenin loci on chromosome arms 1AS, 1BS, and 6DS, with the synthetic hexaploid contributing favorable alleles.Some RI lines showed quality values consistently superior to those of the parental material, suggesting the potential of further evaluating new combinations of alleles from diploid and tetraploid relatives, especially alleles of known storage proteins, for improvement of quality traits in wheat cultivars.

Evaluation of five strategies for obtaining a core subset from a large genetic resource collection of durum wheat

The use of plant genetic resources contained in a large collection may be enhanced by specifying subsamples, called core samples. Five strategies for selecting a core sample from a collection of 3000 durum wheat accessions were applied and evaluated using four qualitative and eight quantitative spike characters. Each of the following strategies generated about 500 accessions for the core sample: random, random-systematic according to chronology of entry of the accessions into the collection, stratified by countryof-origin, stratified by log frequency by country-of-origin, and stratified by canonical variables. The first three strategies produced samples representative of the whole collection, but the remaining two produced the desired effect of increasing frequencies from less-represented countries-of-origin for several characters. The stratified canonical sample increased phenotypic variances. The quality of core samples is dependent upon good passport and evaluation data to partition the collection. The multivariate approach is extremely useful, but requires considerable data from the whole collection. Ecogeographic origin may be used in the absence of evaluation data on several characters to select useful core samples.

Use of recombinant inbred lines of wheat for study of associations of high-molecular-weight glutenin subunit alleles to quantitative traits 1. Grain yield and quality prediction tests

SummaryThe high-molecular-weight glutenin subunits (HMW glutenin), encoded by alleles at homoeologous lociGlu-A1,Glu-B1, andGlu-D1 on the long arms of chromosomes1A,1B, and1D of a set of F8 random recombinant inbred lines (RIL) derived from the bread wheat cross Anza × Cajeme 71, were classified by SDS-PAGE. Anza has poor breadmaking quality and HMW-glutenin subunits (Payne numbers) null (Glu-A1c), 7+8 (Glu-B1b), and 2+12 (Glu-D1a); Cajeme 71 has good quality and 1 (Glu-A1a), 17+18 (Glu-B1i), and 5+10 (Glu-D1d). The combinations of these alleles in the RIL were examined for associations with grain yield and four indicators of grain quality — protein content, yellowberry, pearling index, and SDS sedimentation volume. Data were obtained from a field experiment with three nitrogen fertilization treatments on 48 RIL and the parents. Orthogonal partitioning of the genetic variance associated with the three HMW glutenin subunit loci into additive and epistatic (digenic and trigenic) effects showed strong associations of these loci with grain yield and the indicators of quality; however, the associations accounted for no more than 25% of the differences between the parents. Genetic variance was detected among the RIL, which had the same HMW glutenin genotype for all traits. Epistatic effects were absent for grain yield and yellowberry, but were substantial for grain protein content, pearling index, and SDS sedimentation volume. All three loci had large single-locus additive effects for grain yield, protein, and SDS sedimentation volume. Yellowberry was largely influenced byGlu-B1 andGlu-D1, whereas pearling index was associated withGlu-A1 andGlu-B1. Even though the observed associations-of effects of HMW glutenin loci with the quantitative characters were small relative to the total genetic variability, they are of considerable importance in understanding the genetics of wheat quality, and are useful in the development of new wheat varieties with specific desired characteristics.

Reconstruction of the Synthetic W7984 × Opata M85 wheat reference population

Reference populations are valuable resources in genetics studies for determining marker order, marker selection, trait mapping, construction of large-insert libraries, cross-referencing marker platforms, and genome sequencing. Reference populations can be propagated indefinitely, they are polymorphic and have normal segregation. Described are two new reference populations who share the same parents of the original wheat reference population Synthetic W7984 (Altar84/ Aegilops tauschii (219) CIGM86.940) x Opata M85, an F(1)-derived doubled haploid population (SynOpDH) of 215 inbred lines and a recombinant inbred population (SynOpRIL) of 2039 F(6) lines derived by single-plant self-pollinations. A linkage map was constructed for the SynOpDH population using 1446 markers. In addition, a core set of 42 SSR markers was genotyped on SynOpRIL. A new approach to identifying a core set of markers used a step-wise selection protocol based on polymorphism, uniform chromosome distribution, and reliability to create nested sets starting with one marker per chromosome, followed by two, four, and six. It is suggested that researchers use these markers as anchors for all future mapping projects to facilitate cross-referencing markers and chromosome locations. To enhance this public resource, researchers are strongly urged to validate line identities and deposit their data in GrainGenes so that others can benefit from the accumulated information.

Use of recombinant inbred lines of wheat for study of associations of high-molecular-weight glutenin subunit alleles to quantitative traits : 2. Milling and bread-baking qualitiy.

SummaryRecombinant inbred lines (RILs) derived by single plant descent to F8 from a hybrid of Anza, a low-quality cultivar, and Cajeme 71, a high-quality cultivar, differed in alleles at three high-molecular-weight glutenin (HMW-glu) seed storage protein loci. The 48 RILs were classified by SDS-PAGE for the Anza alleles Glu-Alc (null), Glu-B1b (subunits 7 + 8), and Glu-D1a (subunits 2 + 12) and for Cajeme 71 alleles Glu-A1a (sub-unit 1), Glu-B1I (subunits 17 + 18), and Glu-D1d (subunits 5 + 10). All RILs and parents were grown in a replicated field trial with three levels of nitrogen (N) fertilization. Additive and additive x additive gene effects for the three loci were detected by orthogonal comparisons of means for each of six wheat end-use quality traits. Each HMW-glu genotype was represented by three to ten RILs so that variability among RILs within each HMW-glu genotype could be examined. N effects were consistently small. All traits except flour yield were highly correlated with predictor traits studied earlier. Flour protein content, baking water absorption, dough mixing time, bread loaf volume, and bread loaf crumb score were all correlated, suggesting similar gene control for these traits; however, specific additive locus contributions were evident: αB for flour yield; αB and αD for flour protein; and αB for absorption, but differing in sign; all three loci for mixing time, but αB was negative; and all three loci were positively associated with loaf volume. Digenic epistatic effects were significant for flour yield (αAD), flour protein (αAB), and absorption and mixing time (αAD, αBD). Only flour yield showed a trigenic epistatic effect. Six of seven epistatic effects were negative, thus showing how progress in breeding for high quality may be impeded by interaction of genes which, by themselves, have strong positive additive effects. Considerable genetic variance among RILs within a HMW-glu genotype was detected for all traits, and the summation of α effects accounted for a mean of 13% of the parental differences for the six traits examined in this study. Clearly, further resolution of the genetics of wheat quality would be desirable from a plant breeding point of view.

Biodiversity in Agriculture: Domestication, Evolution, and Sustainability

List of contributors Foreword Bruce D. Smith Acknowledgments Introduction. The domestication of plants and animals: ten unanswered questions Paul Gepts, Robert Bettinger, Stephen Brush, Ardeshir Damania, Thomas Famula, Patrick McGuire and Calvin Qualset 1. The local origins of domestication Jared Diamond Part I. Early Steps in Agricultural Domestication: 2. Evolution of agro-ecosystems: biodiversity, origins and differential development David R. Harris 3. From foraging to farming in western and eastern Asia Ofer Bar-Yosef 4. Predomestic cultivation during the late Pleistocene and early Holocene in the Northern Levant George Willcox 5. New archaeobotanical information on plant domestication from macro-remains: tracking the evolution of domestication syndrome traits Dorian Q. Fuller 6. New archaeobotanical information on early cultivation and plant domestication involving microplant remains Dolores R. Piperno 7. How and why did agriculture spread? Peter Bellwood 8. California Indian proto-agriculture: its characterization and legacy M. Kat Anderson and Eric Wohlgemuth Part II. Domestication of Animals and Impacts on Humans: 9. Pathways to animal domestication Melinda A. Zeder 10. Genetics of animal domestication Leif Andersson 11. Genome-wide approaches for the study of dog domestication Bridgett M. vonHoldt, Melissa M. Gray and Robert K. Wayne 12. Malaria and rickets represent selective forces for the convergent evolution of adult lactase persistence Loren Cordain, Matthew S. Hickey and Kami Kim Part III. Issues in Plant Domestication: 13. The dynamics of rice domestication: a balance between gene flow and genetic isolation Susan R. McCouch, Michael J. Kovach, Megan Sweeney, Hui Jiang and Mande Semon 14. Domestication of lima beans: a new look at an old problem M. I. Chacon S., J. R. Motta-Aldana, M. L. Serrano S. and D. G. Debouck 15. Genetic characterization of cassava (Manihot esculenta Crantz) and yam (Dioscorea trifida L.) landraces in swidden agriculture systems in Brazil Elizabeth A. Veasey, Eduardo A. Bressan, Marcos V. B. M. Siqueira, Aline Borges, Jurema R. Queiroz-Silva, Kayo J. C. Pereira, Gustavo H. Recchia and Lin Chau Ming 16. Pigeonpea - from an orphan to a leader in food legumes C. L. Laxmipathi Gowda, K. B. Saxena, R. K. Srivastava, H. D. Upadhyaya and S. N. Silim Part IV. Traditional Management of Biodiversity: 17. Ecological approaches to crop domestication D. B. McKey, M. Elias, B. Pujol and A. Duputie 18. Agrobiodiversity shifts on three continents since Vavilov and Harlan: assessing causes, processes and implications for food security Gary Paul Nabhan, Ken Wilson, Ogonazar Aknazarov, Karim-Aly Kassam, Laurie Monti, David Cavagnaro, Shawn Kelly, Tai Johnson and Ferrell Sekacucu 19. Indigenous peoples conserving, managing, and creating biodiversity Jan Salick 20. Land architecture in the Maya lowlands: implications for sustainability B. L. Turner II and Deborah Lawrence 21. Agrobiodiversity and water resources in agricultural landscape evolution (Andean Valley irrigation, Bolivia, 1986 to 2008) Karl S. Zimmerer Part V. Uses of Biodiversity and New and Future Domestications: 22. Participatory domestication of indigenous fruit and nut trees: new crops for sustainable agriculture in developing countries Roger R. B. Leakey 23. The introduction and dispersal of Vitis vinifera into California: a case study of the interaction of man, plants, economics, and environment James Lapsley 24. Genetic resources of yeast and other micro-organisms Charles W. Bamforth 25. Biodiversity of native bees and crop pollination with emphasis on California Robbin W. Thorp 26. Aquaculture, the next wave of domestication Dennis Hedgecock 27. Genetic sustainability and biodiversity: challenges to the California dairy industry Juan F. Medrano Index.