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

Recent advances in crop research have the potential to accelerate genetic gains in wheat, especially if co-ordinated with a breeding perspective. For example, improving photosynthesis by exploiting natural variation in Rubisco’s catalytic rate or adopting C4 metabolism could raise the baseline for yield potential by 50% or more. However, spike fertility must also be improved to permit full utilization of photosynthetic capacity throughout the crop life cycle and this has several components. While larger radiation use efficiency will increase the total assimilates available for spike growth, thereby increasing the potential for grain number, an optimized phenological pattern will permit the maximum partitioning of the available assimilates to the spikes. Evidence for underutilized photosynthetic capacity during grain filling in elite material suggests unnecessary floret abortion. Therefore, a better understanding of its physiological and genetic basis, including possible signalling in response to photoperiod or growth-limiting resources, may permit floret abortion to be minimized for a more optimal source:sink balance. However, trade-offs in terms of the partitioning of assimilates to competing sinks during spike growth, to improve root anchorage and stem strength, may be necessary to prevent yield losses as a result of lodging. Breeding technologies that can be used to complement conventional approaches include wide crossing with members of the Triticeae tribe to broaden the wheat genepool, and physiological and molecular breeding strategically to combine complementary traits and to identify elite progeny more efficiently.

/pdf/raising-yield-potential-in-wheat-25io6aksxr.pdf

Raising yield potential in wheat

Summary
This review charts the use of the concept of harvest index in crop improvement and physiology, concentrating on the literature from the last 20 years. Evidence from abstract journals indicates that the term has been applied most to small grain cereal crops and pulses, in India, Western Europe and the USA, and that it has been less useful for maize and tuber crops. Standard methods of measuring harvest index, the associated problems of measurement and interpretation, and representative values for a range of world species are reviewed. The values for modern varieties of most intensively-cultivated grain crops fall within the range 0.4 to 0.6. Variation between varieties of the same species is illustrated by trends in the harvest indices of old, outclassed and recent varieties of temperate and mediterranean wheat and barley (compared under uniform conditions); this shows a progressive increase throughout the present century, although improvement has been much slower in Australia and Canada than in the UK. In most cases, the improvement in harvest index has been a consequence of increased grain population density coupled with stable individual grain weight. The high heritability of harvest index is explored by examining its (rather weak) response to variation in environmental factors (fertilisation, population density, application of growth regulators) in the absence of severe stress. A fuller perspective is gained by reviewing aspects of the harvest index of rice, maize and tropical pulses. With rice, attention must be paid to the fact that the adhering lemma and palea (not primarily part of economic yield) can make up 20% of grain weight; and there are important interactions among biomass, grain yield and season length. Maize differs from most small grain crops in that harvest index (in N. American varieties) was already high at the start of this century, and increases in yield potential have been largely the consequence of increased biomass production. The harvest index of many pulse species and varieties tends to be low because selection has been for some yield in all seasons. Extension of the harvest index concept to express the partitioning of mineral nutrients as well as dry matter (e.g. the nitrogen harvest index) has provided a range of responses whose implications for production and breeding remain to be explored. It is concluded that even though the principal cereal crops appear to be approaching the upper limit of harvest index, and future yield gains will have to be sought by increased biomass production, there will still be a need for the concept of harvest index as a tool in interpreting crop response to different environments and climatic change.

Harvest index: a review of its use in plant breeding and crop physiology

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.

/pdf/raising-yield-potential-of-wheat-iii-optimizing-partitioning-32v5cur56c.pdf

Raising yield potential of wheat. III. Optimizing partitioning to grain while maintaining lodging resistance

Four sets of near-isogenic lines carrying different combinations of the alleles Rht-B1b, Rht-D1b and Rht-B1c for gibberellin-insensitive dwarfism in hexaploid wheat (Triticum aestivum L.) were compared with tall controls in a series of yield trials in eastern England and central Germany. In all four varietal backgrounds the effects of Rht-B1b and Rht-D1b were similar (plant height ≈ 86 and 83% of tall controls respectively) and in combination reduced plant height to c. 58%. The Rht-B1c allele caused more severe dwarfism (c. 50%) and, when combined with Rht-D1b, reduced plant height still further to c. 41%.Data from the trials were consistent with a model for height/yield relationships in which the pleiotropic effects of the Rht alleles on yield can be inferred from their primary function: insensitivity to gibberellin limits stem extension growth, decreasing assimilate demand for this organ and diverting it to the developing ear (which is not itself dwarfed). The net balance between the resulting increase in harvest index and the curvilinear relationship observed between plant height and total shoot yield results in optimum grain yields at intermediate plant heights.Yield advantages of shorter plants over tall controls were evident over several trials with mean grain yields ranging from 200 to 760 g m−2. The optimum plant height for yield improvement in different genetic backgrounds was achieved by different Rht alleles according to the background varietal height, such that intrinsically taller genotypes required more potent Rht alleles to achieve maximum potential grain yield.Ear yield components showed increases in grain number due to Rht pleiotropy, from which it is inferred that the number of grains per ear is limited by supply of assimilates pre-anthesis. Increases in grain number were associated with decreases in mean weight per grain which varied according to severity of dwarfism and varietal background, so that the net effect on grain yield per ear was sometimes positive, sometimes negative, and sometimes neutral in different Rht/variety combinations.

https://www.cambridge.org/core/services/aop-cambridge-core/content/view/S0021859696003942

Optimizing wheat grain yield: effects of Rht (gibberellin-insensitive) dwarfing genes

This study of the effect of major Norin 10 dwarfing genes (Rht1, Rht2) on kernel number in spring wheat emphasised three near-isogenic pairs of spring wheat, differing in the presence or absence of both genes. Plants were grown under controlled environment conditions with equal light intensities at their tops and were restricted to main stems by repeated tiller trimming. The dwarf wheats had a higher proportion of shoot dry weight in the spike at anthesis, an effect which field studies suggest is closely associated with their production of more kernels per unit land area. Stems of dwarf genotypes were somewhat shorter as early as 50 days before anthesis (i.e. before floral initiation) but relative differences became especially great commencing at 35 days before anthesis. Apex and spike lengths were never very different. A greater proportion of dry matter was distributed to leaf lamina from 50 days before anthesis until the end of lamina growth in dwarf wheats, while partitioning to stems was lower from 50 days before anthesis onwards. Partitioning to spikes was only higher in the 15 days preceding anthesis. Stem water-soluble carbohydrate (WSC) contents at anthesis were greater in dwarf wheats but maximum spike WSC contents reached at about 15 days before anthesis were lower. Spike morphogenesis including floret production and grain setting did not differ generally, except for a tendency with dwarf wheats for a longer floral initiation to anthesis interval and for more kernels per unit of spike weight at anthesis. The major unique feature of dwarf genotypes, the higher proportion of dry matter partitioned to the spike, appeared to be due to reduced competition from growing stems in dwarf wheats rather than intrinsic differences in the growth potential of their spikes.

Increased Kernel Number in Norin 10-Derived Dwarf Wheat: Evaluation of the Cause

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Increased Kernel Number in Norin 10-Derived Dwarf Wheat: Evaluation of the Cause