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• W2610391446 abstract "POLICY FORUM S CIENCE GALLEY OVERLINE Genome plasticity a key factor in the success of polyploid wheat under domestication Insert Deck Here Jorge Dubcovsky†, Jan Dvorak Wheat was domesticated approxi- mately 10,000 years ago and has since spread worldwide to become one of the major crops. Its adaptability to diverse environments and end-uses is surprising given the diversity bottlenecks expected from recent domestication and polyploid speciation events. Wheat compensates for these bottlenecks by capturing part of the genetic diversity of its progenitors and by generating new diversity at a relatively fast pace. Frequent gene deletions and dis- ruptions generated by a fast replacement rate of repetitive sequences are buffered by the polyploid nature of wheat, resulting in subtle dosage effects on which selection can operate. With 620 million tons produced annually worldwide, wheat provides approximately one fifth of the calories consumed by hu- mans (1). Roughly 95% of the wheat crop is common wheat, used for making bread, cookies, and pastries, whereas the remaining 5% is durum wheat, used for making pasta and other semolina products. Einkorn wheat and other hulled wheats, namely emmer and spelt, are today relic crops of minor eco- nomic significance (2, 3). While einkorn is a diploid species, durum and common wheat are polyploid species that originated by interspecific hybridization of two and three different diploid species, re- spectively (Fig. 1). The success of these do- mesticated polyploid species parallels the success of natural polyploid species, which represent more than 70% of the plant species (reviewed in (4)), and tend to have a more extended geographic distribution than their close diploid relatives (5). Consequently, re- cent advances in wheat genomics may shed light on the genetic causes of the broad adaptability of natural polyploid plant spe- cies. Department of Plant Sciences, University of California, One Shields Avenue, Davis, CA 95616, USA. To whom correspondence should be addressed Wheat domestication The transition from hunting and gathering to agrarian lifestyles in western Asia was a threshold in the evolution of human societies. Domestication of three cereals – einkorn, emmer, and barley – marked the beginning of that process (6). Genetic relationships be- tween wild and domesticated einkorn and emmer suggest that the region west of Diyar- bakir in southeastern Turkey is the most likely site of their domestication (Fig. 2) (7- 9). From this area, the expansion of agricul- ture lead to the dissemination of domesti- cated einkorn (T. monococcum, genomes A m A m ) and domesticated emmer (T. tur- gidum ssp. dicoccon, genomes BBAA) across Asia, Europe, and Africa. Southwest- ern expansion of domesticated emmer culti- vation resulted in sympatry with the southern subpopulation of wild emmer (T. turgidum ssp. dicoccoides, genomes BBAA). Gene ex- changes between the northern domesticated emmer with southern wild emmer popula- tions or with emmer domesticated in the southern region resulted in the formation of a center of emmer diversity in Southern Levant (Fig. 2) (9). The consequence was a subdivi- sion of domesticated emmer into northern and southern subpopulations with an increase in gene diversity in the latter (9). Northeast expansion of domesticated emmer cultivation resulted in sympatry with Aegilops tauschii (genomes DD) and the emergence of hexap- loid common wheat (T. aestivum, genomes BBAADD) (10) within the corridor stretch- ing from Armenia to the southwestern coastal area of the Caspian Sea (11) (Fig. 2). The genetic changes responsible for the suite of traits that differentiate domesticated plants from their wild ancestors are referred to as the “domestication syndrome” (12). In wheat, as in other cereals, a primary compo- nent of this syndrome was the loss of spike shattering, preventing the grains to be scat- tered by the wind and facilitating harvesting (Fig. 1). Abscission scars of einkorn remains from archeological sites in northern Syria www.sciencemag.org SCIENCE and southeastern Turkey revealed a gradual increase of non-shattering einkorn spikes from 9250 to 6500 years BP, a discovery in- terpreted as evidence of a prolonged domes- tication period of cereals (13). The chromo- some locations of the genes controlling shattering in einkorn are unknown, but in emmer wheat it is determined by the Br (brit- tle rachis) loci on chromosomes 3A and 3B (14) (Fig. 1). Another important trait for wheat domes- tication was the loss of tough glumes, con- verting hulled wheat into free-threshing wheat (Fig. 1). The primary genetic determi- nants of the free-threshing habit are recessive mutations at the Tg (tenacious glume) loci (15), accompanied by modifying effects of the dominant mutation at the Q locus and mutations at several other loci (15). The re- cent cloning of Q, which also controls the square spike phenotype in common wheat, showed that it encodes an AP2-like transcrip- tion factor. The mutation that gave rise to the Q allele is the same in tetraploid and hexap- loid free-threshing wheats suggesting that it occurred only once (16). Seeds of free-threshing wheat began to appear in archeological sites about 10,000 years before present (BP) (17). The tetraploid forms of these Neolithic free-threshing wheat may be the ancestor of the modern large- seeded, free-threshing durum (Fig. 1), which is genetically most closely related to the Mediterranean and Ethiopian subpopulations of domesticated emmer (Fig. 2) (9). The first archeological records of durum appeared in Egypt during the Greco-Roman times (re- viewed in (2)). Other traits of the wheat domestication syndrome shared by all domesticated wheats are increased seed size (Fig. 1A-B), reduced number of tillers, more erect growth, and re- duced seed dormancy. One gene affecting seed size is GPC-B1, an early regulator of senescence with pleiotropic effects on grain nutrient content (18). In some genotypes and environments the accelerated grain maturity VOL. xxx • galley printed 21 May, 2007 • • For Issue Date: ????" @default.
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• W2610391446 title "Erratum: Genome plasticity a key factor in the success of polyploid wheat under domestication (Science (1862))" @default.
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