Winter wheat (Triticum aestivum L.) was grown for 4 years in multi-factorial field trials at Rothamsted, southern England. Thirty nine elite commercial cultivars (primarily short-straw) were grown including those released in the UK over a 25-year period, a selection of continental varieties, and three older, tall varieties. Varieties spanned the quality spectrum from 'bread' to 'feed'. The crops were given ammonium nitrate at five rates in the range 0-350 kg-N/ha as a 3-way split. The aim was to quantify the genotypic variation in total nitrogen uptake by grain and straw (total-Nup), and in nitrogen utilization efficiency for grain yield (grain yield per unit of N taken up) (grain-NutE). Depending on treatment, grain yield ranged from 2.1 to 11.8 t/ha (85% DM), grain %N from 1.1% to 2.8% (in DM), total-Nup from 31 to 264 kg-N/ha, and grain-NutE from 27 to 77 kg-DM/kg-N. There were significant varietal differences in total N-uptake and grain-NutE both between 'tall' and 'short' varieties and within 'short' varieties. The best short varieties took up 31-38 kg/ha more N than the worst, and grain-NutE was 24-42% better, depending on N-rate. Up to 77% of the variation in grain-NutE was accounted for by yield. All interactions between the factors 'Variety', 'Year', and 'N-rate' were highly significant, but only 'Year × N-rate' made an important contribution to the variation. There was a near-functional inverse relationship between grain-NutE and grain %N; high-quality wheat (high grain %N) can be expected to have a low grain-NutE. The four key variables determining N-efficiency in a wheat crop-grain yield, grain %N, total N-uptake and nitrogen harvest index (NHI)-are ultimately constrained by the law of conservation of matter. Improving grain-NutE for fixed total-Nup and NHI can only be achieved at the expense of grain %N. To improve grain-NutE and maintain grain %N requires a simultaneous increase in NHI and grain starch yield which may be difficult to achieve in practice. The law of conservation of matter ultimately sets a limit on the physiological and agronomic processes that determine crop N requirements. A high yield of high-quality grain (high grain %N) requires a high input and uptake of nitrogen.

Interactions between sulfur (S) nutritional status and sulfate transporter expression in field-grown wheat (Triticum aestivum) were investigated using Broadbalk +S and 2S treatments (S fertilizer withheld) at Rothamsted, United Kingdom. In 2008, S, sulfate, selenium (Se), and molybdenum (Mo) concentrations and sulfate transporter gene expression were analyzed throughout development. Total S concentrations were lower in all tissues of 2S plants, principally as a result of decreased sulfate pools. S, Se, and Mo concentrations increased in vegetative tissues until anthesis, and thereafter, with the exception of Mo, decreased until maturity. At maturity, most of the S and Se were localized in the grain, indicating efficient remobilization from vegetative tissues, whereas less Mo was remobilized. At maturity, Se and Mo were enhanced 7-and 3.7-fold, respectively, in 2S compared with +S grain, while grain total S was not significantly reduced. Enhanced expression of sulfate transporters, for example Sultr1;1 and Sultr4;1, in 2S plants explains the much increased accumulation of Se and Mo (7-and 3.7-fold compared with +S in grain, respectively). Sultr5;2 (mot1), thought to be involved in Mo accumulation in Arabidopsis (Arabidopsis thaliana), did not fully explain patterns of Mo distribution; it was expressed in all tissues, decreasing in leaf and increasing in roots under 2S conditions, and was expressed in florets at anthesis but not in grain at any other time. In conclusion, S fertilizer application has a marked impact on Mo and Se distribution and accumulation, which is at least partially a result of altered gene expression of the sulfate transporter family.

2-carboxy-D-arabinitoI-l-phosphate (CAIP) bound toRubisco eitber in leaf extracts or after purification can be displaced by SO4^ ions. Tbus, treatment of leaf extracts witb a buffer containing 200 mol m"^d isplaces any 4 p y bound CAIP and enables measurement of maximum carboxylation potential. In tobacco leaves, tbe activity following treatment witb SO4^ ions ('maximal activity') is greater tban tbe total Rubisco activity. Tbe ratio of tbe two activities altered in a dynamic way witb fluctuations in irradiance. Even in species wbicb do not produce significant amounts of CAIP, tbe maximal activity greatly exceeded tbe total activity. Anion excbange separation of components in acid extracts confirmed tbe absence of CAIP in tobacco leaves barvested above an irradiance of 300 //mol quanta m"^ s~*, but tbe presence of anotber inbibitor of Rubisco. Tbese results are consistent witb tbe regulation of Rubisco activity by inbibitors otber tban CAIP wbicb, like CAIP, can be displaced by SO4^~ ions.

Increasing demands for productivity together with environmental concerns about fertilizer use dictate that the future sustainability of agricultural systems will depend on improving fertilizer use efficiency. Characterization of the biological processes responsible for efficient fertilizer use will provide tools for crop improvement under reduced inputs. Transcriptomic and metabolomic approaches were used to study the impact of nitrogen (N) and sulphur (S) deficiency on N and S remobilization from senescing canopy tissues during grain filling in winter wheat (Triticum aestivum). Canopy tissue N was remobilized effectively to the grain after anthesis. S was less readily remobilized. Nuclear magnetic resonance (NMR) metabolite profiling revealed significant effects of suboptimal N or S supply in leaves but not in developing grain. Analysis of amino acid pools in the grain and leaves revealed a strategy whereby amino acid biosynthesis switches to the production of glutamine during grain filling. Glutamine accumulated in the first 7 d of grain development, prior to conversion to other amino acids and protein in the subsequent 21 d. Transcriptome analysis indicated that a down-regulation of the terminal steps in many amino acid biosynthetic pathways occurs to control pools of amino acids during leaf senescence. Grain N and S contents increased in parallel after anthesis and were not significantly affected by S deficiency, despite a suboptimal N:S ratio at final harvest. N deficiency resulted in much slower accumulation of grain N and S and lower final concentrations, indicating that vegetative tissue N has a greater control of the timing and extent of nutrient remobilization than S.