A substantial increase in grain yield potential is required, along with better use of water and fertilizer, to ensure food security and environmental protection in future decades. For improvements in photosynthetic capacity to result in additional wheat yield, extra assimilates must be partitioned to developing spikes and grains and/or potential grain weight increased to accommodate the extra assimilates. At the same time, improvement in dry matter partitioning to spikes should ensure that it does not increase stem or root lodging. It is therefore crucial that improvements in structural and reproductive aspects of growth accompany increases in photosynthesis to enhance the net agronomic benefits of genetic modifications. In this article, six complementary approaches are proposed, namely: (i) optimizing developmental pattern to maximize spike fertility and grain number, (ii) optimizing spike growth to maximize grain number and dry matter harvest index, (iii) improving spike fertility through desensitizing floret abortion to environmental cues, (iv) improving potential grain size and grain filling, and (v) improving lodging resistance. Since many of the traits tackled in these approaches interact strongly, an integrative modelling approach is also proposed, to (vi) identify any trade-offs between key traits, hence to define target ideotypes in quantitative terms. The potential for genetic dissection of key traits via quantitative trait loci analysis is discussed for the efficient deployment of existing variation in breeding programmes. These proposals should maximize returns in food production from investments in increased crop biomass by increasing spike fertility, grain number per unit area and harvest index whilst optimizing the trade-offs with potential grain weight and lodging resistance.

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Society relies heavily on inorganic phosphorus (P) compounds throughout its food chain. This dependency is not only very inefficient and increasingly costly but is depleting finite global reserves of rock phosphate. It has also left a legacy of P accumulation in soils, sediments and wastes that is leaking into our surface waters and contributing to widespread eutrophication. We argue for a new, more precise but more challenging paradigm in P fertilizer management that seeks to develop more sustainable food chains that maintain P availability to crops and livestock but with reduced amounts of imported mineral P and improved soil function. This new strategy requires greater public awareness of the environmental consequences of dietary choice, better understanding of soil-plant-animal P dynamics, increased recovery of both used P and unutilized legacy soil P, and new innovative technologies to improve fertilizer P recovery. In combination, they are expected to deliver significant economic, environmental, and resource-protection gains, and contribute to future global P stewardship.

The efficient use of fertilizer nitrogen (N) is crucial to sustainable human nutrition. All crops receive significant amounts of additional N in temperate environments, through fixation or fertilizer use. This paper reviews progress towards the efficient use of fertilizer N by winter wheat (Triticum aesitivum L.) and spring barley (Hordeum vulgare L.) in the UK, acknowledging that on-farm this is governed by economics. Recent multi-site N response experiments on old and modern varieties show that yield improvements since the 1980s have been accompanied by increases in economic optimum N amounts for wheat but not for spring barley. On-farm N use efficiency (NUE) has increased for barley because increased yields with optimum N were associated with compensatory decreases in grain N concentration, whereas on-farm NUE has not increased for wheat because grain N concentration has not changed and improvements in N capture were insufficient to make up for the increased yield. Genetic effects on NUE are shown to differ markedly depending on whether they are determined at a single N rate, as in variety trials, or with optimum N amounts. It is suggested that, in order to elicit faster improvement in NUE on farms, breeding and variety testing should be conducted at some sites with more than one level of applied N, and that grain N%, N harvest index, and perhaps canopy N ratio (kg N ha(-1) green area) should be measured more widely. It is also suggested that, instead of using empirical functions, N responses might be analysed more effectively using functions based on explanations of yield determination for which the parameters have some physiological meaning.

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A quantitative model of wheat root systems is developed that links the size and distribution of the root system to the capture of water and nitrogen (which are assumed to be evenly distributed with depth) during grain filling, and allows estimates of the economic consequences of this capture to be assessed. A particular feature of the model is its use of summarizing concepts, and reliance on only the minimum number of parameters (each with a clear biological meaning). The model is then used to provide an economic sensitivity analysis of possible target characteristics for manipulating root systems. These characteristics were: root distribution with depth, proportional dry matter partitioning to roots, resource capture coefficients, shoot dry weight at anthesis, specific root weight and water use efficiency. From the current estimates of parameters it is concluded that a larger investment by the crop in fine roots at depth in the soil, and less proliferation of roots in surface layers, would improve yields by accessing extra resources. The economic return on investment in roots for water capture was twice that of the same amount invested for nitrogen capture.

The plant characteristics required to make wheat lodging-proof for the least investment of biomass are calculated using a validated model of the lodging process and information about the dry matter costs of altering lodgingassociated characters. The plant characteristics required to give a crop yielding 8 t ha -1 with 500 shoots m -2 and 200 plants m -2 , a lodging return period of 25 years include a height of 0.7 m, a root plate spread of 57 mm, and for the bottom internode a wall width of 0.65 mm with a diameter/material strength combination ranging from a diameter of 5.86 mm with a material strength of 20 MPa to a diameter of 4.00 mm with a material strength of 50 MPa. It is estimated that this ideotype would require 7.9 t ha -1 of stem biomass and would have a harvest index of 0.42. Observations of a wide range of varieties grown using crop management to maximise lodging resistance without reducing yield potential showed that the root plate of the best variety was 7 mm less than the ideotype target, the stem character targets were achieved but not all in one variety, and the height target was achievable with the use of plant growth regulators. Plant breeders must therefore focus on selecting for a wider root plate and combining the appropriate stem strength characteristics.

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