Waiting for fine times: genetics of flowering time in wheat

To maximise yield potential in any environment, wheat cultivars musthave an appropriate flowering time and life cycle duration which`fine-tunes' the life cycle to the target environment. This in turn, requiresa detailed knowledge of the genetical control of the key components of thelife cycle. This paper discusses our current knowledge of the geneticalcontrol of the three key groups of genes controlling life-cycle duration inwheat, namely those controlling vernalization response, photoperiodresponse and developmental rate (`earliness per se', Eps genes).It also discusses how our ability to carry out comparative mapping of thesegenes across Triticeae species, and particularly with barley, is indicatingnew target genes for discovery in wheat. Major genes controllingvernalization response, the Vrn-1 series, have now been located bothgenetically and physically on the long arms of the homoeologous group fivechromosomes. These genes are homoeologous to each other and to thevernalization genes on chromosomes 5H of barley and 5R of rye.Comparative analysis with barley also indicates that other series ofvernalization response genes may exit on chromosomes of homoeologousgroups 4 (4B, 4D, 5A) and 1. The major genes controlling photoperiodresponse in wheat, the Ppd-1 genes, are located on the homoeologousgroup 2 chromosomes, and are homoeologous to a gene on barleychromosome 2H. Mapping in barley also indicates a photoperiod responselocus on barley 1H and 6H, indicating that a homoeologous series shouldexist on wheat group 1 and 6 chromosomes. In wheat, only a few`earliness per se loci have been located, such as on chromosomes ofhomoeologous group 2. However, in barley, all chromosomes appear tocarry such loci, indicating that several series of loci that affectdevelopmental rate independent of environment remain to be discovered.Overall, comparative studies indicate that there are probably twenty-fiveloci controlling the duration of the life-cycle, Vrn, Ppd and Eps genes, that remain to be mapped in wheat. There are major gaps inour knowledge of the detailed physiological effects of genes discovered todate on the timing of the life cycle from different sowing dates. This isbeing addressed by studying the phenology of isogenic and deletion lines inboth field and controlled environmental conditions. This has indicated thatthe vernalization genes have major effects on the rate of primodiaproduction, whilst the photoperiod genes affect the timing of terminalspikelet production and stem elongation, and these effects interact withsowing date.