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 telomeretelomere 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.dysploidy ͉ linkage map ͉ rice ͉ sorghum ͉ wheat

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.

Yield in cereals is a function of seed number and weight; both parameters are largely controlled by seed sink strength. The allosteric enzyme ADP-glucose pyrophosphorylase (AGP) plays a key role in regulating starch biosynthesis in cereal seeds and is likely the most important determinant of seed sink strength. Plant AGPs are heterotetrameric, consisting of two large and two small subunits. We transformed wheat (Triticum aestivum L.) with a modified form of the maize (Zea mays L.) Shrunken2 gene (Sh2r6hs), which encodes an altered AGP large subunit. The altered large subunit gives rise to a maize AGP heterotetramer with decreased sensitivity to its negative allosteric effector, orthophosphate, and more stable interactions between large and small subunits. The Sh2r6hs transgene was still functional after five generations in wheat. Developing seeds from Sh2r6hs transgenic wheat exhibited increased AGP activity in the presence of a range of orthophosphate concentrations in vitro. Transgenic Sh2r6hs wheat lines produced on average 38% more seed weight per plant. Total plant biomass was increased by 31% in Sh2r6hs plants. Results indicate increased availability and utilization of resources in response to enhanced seed sink strength, increasing seed yield, and total plant biomass. W heat (Triticum aestivum L.) is one of the world's most important crop plants. An estimated 610 million metric tons of wheat seed were harvested worldwide in the 1997͞98 growing season (1). Wheat seed yield is determined by seed number and weight. Starch is the major component of wheat yield, comprising 70% of wheat seed dry weight. ADP-glucose pyrophosphorylase (AGP; EC 2.7.7.27), an allosterically regulated heterotetramer consisting of two large and two small subunits, catalyzes the rate-limiting reaction in starch biosynthesis in plants (reviewed in refs. 2 and 3). AGP uses the substrates glucose 1-phosphate and ATP to produce ADPglucose and pyrophosphate (4). ADP-glucose is then used as the glucose donor for starch synthases. The positive allosteric effector of AGP is 3-phosphoglycerate, whereas the negative allosteric effector is orthophosphate (Pi) (2, 3). Maize endosperm AGP (5) large subunits are encoded by Shrunken2 (Sh2) (6), whereas small subunits are encoded by Brittle2 (Bt2) (7).Inhibition of AGP activity by Pi appears to limit starch biosynthesis and yield in crop plants. Evidence for this has come from studies of a modified AGP large subunit in maize (8) and from use of an altered bacterial AGP in potato (9). In both cases, AGP modification caused reduced sensitivity to Pi inhibition, resulting in enhanced yield, manifested as increased seed weight in maize (8) and increased starch content in potato (9). Instability of AGP may be an additional limitation on cereal yield. AGP is perhaps the most heat-labile starch biosynthetic enzyme in maize (10-12). A single amino acid substitution in the maize large subunit that conditions more stable large subunit-small subunit interactions (13) may lead to enhanced AGP activity and increa...