The need to accelerate breeding for increased yield potential and better adaptation to drought and other abiotic stresses is an issue of increasing urgency. As the population continues to grow rapidly, the pressure on resources (mainly untouched land and water) is also increasing, and potential climate change poses further challenges. We discuss ways to improve the efficiency of crop breeding through a better physiological understanding by both conventional and molecular methods. Thus the review highlights the physiological basis of crop yield and its response to stresses, with special emphasis on drought. This is not just because physiology forms the basis of proper phenotyping, one of the pillars of breeding, but because a full understanding of physiology is also needed, for example, to design the traits targeted by molecular breeding approaches such as marker-assisted selection (MAS) or plant transformation or the way these traits are evaluated. Most of the information in this review deals with cereals...

Breeding for Yield Potential and Stress Adaptation in Cereals

Continuous increase in the yield of maize (Zea mays L.) in the U.S. Corn Belt has involved an interaction with plant density. A number of contributing traits and mechanisms have been suggested. In this study we used a modeling approach to examine whether changes in canopy and/or root system architecture might explain the observed trends. A maize crop model was generalized so that changes in canopy and root system architecture could be examined. A layered, diurnal canopy photosynthesis model was introduced to predict consequences of change in canopy architecture. A two-dimensional root exploration model was introduced to predict consequences of change in root system architecture. Field experiments were conducted to derive model parameters for the base hybrid (Pioneer 3394). Simulation studies for various canopy and root system architectures were undertaken for a range of sites, soils, and densities. Simulated responses to density compared well with those found in field experiments. The analysis indicated that (i) change in root system architecture and water capture had a direct effect on biomass accumulation and historical yield trends; and (ii) change in canopy architecture had little direct effect but likely had important indirect effects via leaf area retention and partitioning of carbohydrate to the ear. The study provided plausible explanations and identified testable hypotheses for future research and crop improvement effort.

Can Changes in Canopy and/or Root System Architecture Explain Historical Maize Yield Trends in the U.S. Corn Belt?

Bioenergy from plants, particularly from perennial grasses and trees, could make a substantial contribution to alleviation of global problems in climate change and energy security if high yields can be sustained. Here, yield traits in a range of key bioenergy crops are reviewed, from which several targets for future improvement can be identified. Some are already the focus of genetically modified (GM) and non-GM approaches. However, the efficient growth strategies of perennial bioenergy crops rely on newly assimilated and recycled carbon and remobilized nitrogen in a continually shifting balance between sources and sinks. This balance is affected by biotic (e.g. pest, disease) and abiotic (e.g. drought) stresses. Future research should focus on three main challenges: changing (photo)thermal time sensitivity to lengthen the growing season without risking frost damage or limiting remobilization of nutritional elements following senescence; increasing aboveground biomass without depleting belowground reserves required for next year's growth and thus without increasing the requirement for nutrient applications; and increasing aboveground biomass without increasing water use.

Bioenergy from plants and the sustainable yield challenge

Improving crops yield under water-limited conditions is the most daunting challenge faced by breeders. To this end, accurate, relevant phenotyping plays an increasingly pivotal role for the selection of drought-resilient genotypes and, more in general, for a meaningful dissection of the quantitative genetic landscape that underscores the adaptive response of crops to drought. A major and universally recognized obstacle to a more effective translation of the results produced by drought-related studies into improved cultivars is the difficulty in properly phenotyping in a high-throughput fashion in order to identify the quantitative trait loci that govern yield and related traits across different water regimes. This review provides basic principles and a broad set of references useful for the management of phenotyping practices for the study and genetic dissection of drought tolerance and, ultimately, for the release of drought-tolerant cultivars.

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Phenotyping for drought tolerance of crops in the genomics era.

Drought is a predominant cause of low yields worldwide. There is an urgent need for more water efficient cropping systems facing large water consumption of irrigated agriculture and high unproductive losses via runoff and evaporation. Identification of yield-limiting constraints in the plant–soil–atmosphere continuum are the key to improved management of plant water stress. Crop ecology provides a systematic approach for this purpose integrating soil hydrology and plant physiology into the context of crop production. We review main climate, soil and plant properties and processes that determine yield in different water-limited environments. From this analysis, management measures for cropping systems under specific drought conditions are derived. Major findings from literature analysis are as follows. (1) Unproductive water losses such as evaporation and runoff increase from continental in-season rainfall climates to storage-dependent winter rainfall climates. Highest losses occur under tropical residual moisture regimes with short intense rainy season. (2) Sites with a climatic dry season require adaptation via phenology and water saving to ensure stable yields. Intermittent droughts can be buffered via the root system, which is still largely underutilised for better stress resistance. (3) At short-term better management options such as mulching and date of seeding allow to adjust cropping systems to site constraints. Adapted cultivars can improve the synchronisation between crop water demand and soil supply. At long term, soil hydraulic and plant physiological constraints can be overcome by changing tillage systems and breeding new varieties with higher stress resistance. (4) Interactions between plant and soil, particularly in the rhizosphere, are a way towards better crop water supply. Targeted management of such plant–soil interactions is still at infancy. We conclude that understanding site-specific stress hydrology is imperative to select the most efficient measures to mitigate stress. Major progress in future can be expected from crop ecology focussing on the management of complex plant (root)–soil interactions.

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Management of crop water under drought: a review

Understanding the changes underlying past breeding progress may help to focus research efforts and accelerate future genetic gains. The major abiotic stress affecting maize production on a worldwide basis is drought. We addressed the improvements in drought tol- erance over a 50-year period of hybrid breeding by evalu- ating, under targeted stress conditions, a set of 18 Pio- neer-brand hybrids that had been released during the 1953-2001 period. Stress treatments were designed as overlapping windows of water deficit covering the pre- flowering to late grain filling development stages. Data were collected on grain yield, yield components and an- thesis-silking interval (ASI), and were analyzed using a linear mixed model approach. Genetic gain was mea- sured as the slope of the regression of the trait on the year of hybrid release. Significant, positive genetic gains of varying magnitude were observed for grain yield in all windows of stress evaluated. The largest genetic gains for grain yield were observed under conditions of full irriga- tion and severe flowering stress. ASI and barrenness, es- pecially under stress at flowering, were significantly re- duced by selection. Though flowering remains the most susceptible stage to drought in maize, selection has re- duced its negative effects, and susceptibility during early grain filling is now of similar importance in many modern hybrids. Yield under drought at flowering has more than kept pace with the increase in yield potential because of the emphasis breeders have placed on improved floral synchrony.

H. Campos

Papers

Changes in drought tolerance in maize associated with fifty years of breeding for yield in the US Corn Belt [Zea mays L.]