Triticum (Aegilops) tauschii in the natural and artificial synthesis of hexaploid wheat

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Abstract
An account is given of the possible time(s) and place(s) of the origin of hexaploid wheat from natural hybridisation between Triticum tauschii (Ae. tauschii) and both wild and cultivated forms of tetraploid wheat. A recapitulation is presented of the likely genotypic and phenotypic status of the newly arisen natural hexaploid and the likely path of hybridisation from whence it arose. Recent substantial contributions of T. tauschii to wheat improvement indicate the likelihood that introgession en masse from T. tauschii has not occurred throughout its natural and agricultural associations with wheat. This has been substantiated in comparative studies revealing higher levels of genetic variation in T. tauschii compared with the D genome of hexaploid wheat. A case is made for a widening of the concept of the gene pool of T. tauschii for wheat improvement and the notion of a secondary gene pool is proposed to include variation in T. tauschii as it occurs in several polyploid forms of ‘grass Triticum’. The likely differentiation of growth habit forms, conditioned by vernalisation (i.e. vrn) genes, in hexaploid wheat synthesis, including the interaction of these genes in hexaploid wheat, is discussed. It is speculated that growth habit differentiation was of significance to the hexaploid’s yield contribution and survival in tetraploid-hexaploid mixtures (likely to be a common constitution of wheat crops of early agriculture), and in the Neolithic spread of agriculture to the higher latitude, and colder environments of NW Europe and central Asia. The significance of the contribution of T. tauschii to the unique milling and bread-making properties of hexaploid wheat is discussed in the light of Roman discernment of its closer fulfilment of the requirements of leavened bread-making compared with tetraploid wheat. The significance of the contribution of T. tauschii to the evolution of wheat appears to have been much delayed (by ~6500 years) in that hexaploid wheat did not receive singular attention and cultivation until during the Roman era, from whence it gradually rose in popularity to eventually achieve its current pre-eminent status. Continuing systematic evaluation of genetic variation in both the primary and secondary gene pools of T. tauschii for wheat improvement, using both conventional and genetic analysis and contemporary genomic tools, is advocated. The latter approach is particularly important for quantitative traits in the light of wide divergence in plant phenotype of their representatives from that of hexaploid wheat.