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Euphytica (2007) 157:281285 DOI 10.



Challenges to international wheat breeding

Matthew P. Reynolds Hans-Joachim Braun Julian Pietragalla Rodomiro Ortiz

Received: 25 June 2007 / Accepted: 9 July 2007 / Published online: 29 August 2007 Springer Science+Business Media B.V. 2007

This special issue of Euphytica contains selected articles from talks given at the global symposium Challenges to International Wheat Improvement that was organized by the International Maize and Wheat Improvement Center (CIMMYT), in March 2006 at Obregon, Mexico, with support from the Australian Centre for International Agricultural Research (ACIAR). The aim of this symposium was to bring together wheat researchers worldwide to present and discuss their ideas on how to address some of these pressing issues on how to increase wheat production in a sustainable manner; 160 scientists attended from over 30 wheat producing countries. The Journal of Agricultural Science, Cambridge has included some of the articles dealing with physiological basis of yield potential and new technologies for increasing input use eYciency of wheat-based cropping systems agronomy (see Reynolds et al. 2007a), whereas this special issue of Euphytica includes the remaining 17

M. P. Reynolds (&) H.-J. Braun J. Pietragalla R. Ortiz International Maize and Wheat Improvement Center (CIMMYT), Mexico, DF 06600, Mexico e-mail: m.reynolds@cgiar.org H.-J. Braun e-mail: h.braun@cgiar.org J. Pietragalla e-mail: jpietra@gmail.com R. Ortiz e-mail: r.ortiz@cgiar.org

selected articles dealing solely with wheat genetic improvement. The symposium was opened with an address by Nobel Lauretae Dr. Normal Borlaug entitled Sixtytwo years of Wghting hunger: personal recollections. He describes the evolution of international wheat breeding including how shuttle breeding was adopted in Mexico enabling photoperiod sensitivity to be overcome, a pivotal step in creating internationally adapted germplasm. His talk touches on a number of historical yet topical issues, including how the 15b stem rust epidemic in the US in the 1950s is being mirrored 50 years later with the virulent new race Ug99 from East Africa. The evolution of internationally coordinated public goods research in agriculture which led to the formation of the CGIAR (Consultative Group of International Agricultural Research); a system which despite its humanitarian mandate and many successes suVers from declining investment eroding the promise of food security for many of the worlds most resource-poor people. He speciWcally addresses high yield agriculture and the environment, agro-forestry, drought tolerance, the promise of biotechnology, bureaucracies and fear of change, and Wnishes with a quote from 1949 Nobel peace prize winner Lord John Boyd Orr, World Peace Will Not Be Built on Empty Stomachs. Sorrels (2007) reviewed impacts of new technologies in his article Application of new knowledge, technologies, and strategies to wheat improvement. He highlights the complexity of the genomes of



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graminaceous crops and the fact that they are rapidly evolving and heterogeneous, even within species. Not surprisingly the use of marker-assisted selection for improving complex traits remains one of the challenges facing wheat breeders. Progress in recent years includes new transformation protocols, statistical methods, methods for characterizing environments, and equipment for phenotyping traits. He also mentions progress in the area of molecular markers and microarrays applications, gene silencing protocols, DNA sequencing, and transgenic crops. Comparative mapping and QTL studies have provided information about the location, identity and number of genes controlling some economically important traits. The articles by William et al. and Ogbonnaya et al. review advances at the frontiers of wheat improvement research: namely the use of molecular breeding tools and wild species for re-synthesizing wheat, respectively. William et al. argue that markers are being used now to better characterize parental lines improving the eVectiveness of crossing strategies, as well as for tracking genes in segregating progenies. Although still costly, marker-aided selection (MAS) appears to be routinely used for a few traits by wheat breeding programs worldwide. The genetic potential of re-synthesized hexaploid germplasm (when crossed to elite cultivars) was investigated by Ogbonnaya et al. They found that such synthetic-derived lines yielded 8% to 30% higher than the best local check in multi-site trials across diverse regions of Australia. Their results reinforce previous research conducted at CIMMYT that lines derived from synthetic wheat have the potential to signiWcantly improve grain yield across environments. In this regard, Brennan and Martins article Returns to investment in new breeding technologies advocates that breeding programs should carefully assess the likely economic returns from value of incorporating new approaches into their programs, decision likely being based on the scale of the breeding operation, with low cost investments being more universally accessible. In order to meet future demands for wheat, all available technologies must achieve an annual yield increase of about 2% p.a. until 2020. Singh et al. in their article High yielding spring bread wheat germplasm for global irrigated and rainfed production systems report that grain yields of the best new entries were 10% higher than the local checks in international yield trials. While not all genotypes

respond as well across sites, analysis of genotype environment interaction provides opportunities to select for stable genotypes. As outlined by Ortiz et al., international wheat improvement at CIMMYT has included shuttle breeding at two contrasting locations in Mexico to facilitate selection of genotypes with wide adaptation, durable rust and Septoria resistances while incorporating a wide a range of genetic diversity as possible into the thousand or so new entries that are distributed as international nurseries annually. Their article points out that CIMMYTs primary generic target product has been genetically enhanced seed-embedded technology which considers both strategic germplasm enhancement and adaptive breeding to megaenvironments. It is discussed whether CIMMYT and similar CGIAR centers will in the future invest more resources in strategic germplasm enhancement while adaptive breeding would be conducted progressively more by National Agricultural Research Systems (NARS). Strategic germplasm enhancement would include identiWcation and utilization of novel genetic variation, e.g. from landraces and wild speciesincluding production of re-synthesized wheat. Given the mission of IARCs within the international development assistance community, if their products are to change signiWcantly they must consider the needs and relative strength of NARS on a case by case basis. The article by Trethowan and Crossa analyzed 40 years of international spring bread wheat trials. The analysis conWrmed the relevance of the shuttle breeding between two locations in Mexico for global wheat improvement, since selection environments generated in Mexico associate well with global target areas. They describe how integrating information from international sites with that obtained in Mexico helps to improve the eYciency of CIMMYTs global wheat breeding eVort. For more than 40 years, cooperating breeders from many countries have grown these trials, provided their elite germplasm, and returned data to CIMMYT, which made the compiled results available to all cooperators. Without this unprecedented global cooperation, none of the impacts, for example in improving yield under favorable and marginal environments as well as disease resistance would have been possible (Reynolds and Borlaug 2007). Chapman et al. and Ortiz-Ferrara et al. articles assess the advantages of using a global approach by incorporating key genes (e.g. for plant


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height) in wheat breeding lines and emphasizing regional eVorts through participatory research and client-oriented plant breeding, respectively. In the article Relationships between height and yield in near-isogenic spring wheats that contrast for major reduced height genes Chapman and co-authors showed how the environment inXuenced the phenotypic eVects of two major dwarf genes (Rht1 and Rht2). Their results conWrm the advantage of incorporating such genes in wheat cultivars since there was a ca. 10% yield gain for the lines possessing such genes, which was more evident in trials on which the mean height of semi-dwarf isolines exceeded about 80 cm. Ortiz-Ferrara et al. describe in their article Partnering with farmers to accelerate adoption of new technologies in South Asia to improve wheat productivity how several farmer-preferred technologies have been identiWed for adverse conditions in eastern India and Nepal. Due to this participatory-research approach, grain harvests by resource-poor farmers signiWcantly increased (1570%) in locations where farmers, scientists, extension specialists, non-governmental organizations and the private sector engaged in participatory varietal selection. Advances in wheat improvement must also considered wheat Wnal end-use. Negative correlations between grain yield, grain protein concentration, and Wnal end-use are described by De Pauw et al. in the article Shifting undesirable correlations. They concluded that the undesirable correlations of grain yield, grain protein concentration, and time to maturity can be shifted by developing plants, which eYciently produce and partition carbohydrates to grain yield and have improved nitrogen- and water-use eYciency. Improvements in these traits could also be transferred to wheat cultivars in water and nitrogen deWcient areas. They showed that simultaneous selection for quantitative and quality traits with the inclusion of marker assisted selection can shift these undesirable correlations. Duveiller and coauthors present strategies aimed at minimizing or controlling yield losses from major diseases and pests relevant to intensive irrigated wheat systems in the developing world. Options suggested include integrated crop management practices, breeding for genetic resistance, rotations, minimizing physiological stresses and consequent susceptibility by timely sowing and adequate use of fertilizers, and

fungicide application. In their article The challenges of maintaining wheat productivity: pests, diseases, and potential epidemics they also advise about the risk of changes in disease spectrums as a result of climate changes and demonstrate the complex relationships among crop physiology, disease resistance and yield. This special issue of Euphytica also includes an overall assessment of wheat improvement in major grain baskets of the world: South Asia and China. Joshi and co-workers point out that India faces a critical challenge in maintaining food security in the face of its growing population. Indian wheat breeders should therefore aim to improve the crop to address heat stress (exacerbated by global warming due to climate change), water scarcity due dwindling water supplies for irrigation, a growing threat of new virulence of diseases such as wheat rusts and leaf blight, continuous adoption of zero-till and other resource conservation technologies particularly in the intensive and highly productive rice-wheat systems and a high demand for better quality wheat. Challenges to Wheat Production in South Asia in terms of biotic and abiotic stresses are also described by Chatrath et al. He points to the stagnating wheat yields and the declining productivity of wheatrice systems due to intensive tillage and burning of residues leading to the depletion of soil organic carbon. Excessive nutrient mining, imbalanced fertilization and over-exploitation of water resources are the other factors responsible for declining productivity. Addition of organic matter to soil through green manuring and crop residue recycling, balanced fertilization, integrated nutrient management, and crop diversiWcation are suggested to improve total productivity in the regionsee also Gupta and Sayre (2007) for their analysis of the beneWts of conservation agriculture in the region. IdentiWcation of wheat genotypes with high and stable grain yield is of particular relevance for poor farmers. Sharma et al. report results for the Eastern Gangetic Plains Yield Trials, grown in India, Nepal and Bangladesh for the years 1999 to 2005. Lines with improved yield stability and disease resistance were identiWed and released, which underlines the importance and relevance of regional wheat breeding programs. Similarly, Zhou et al. evaluated genetic gains for grain yield in two regions of the Southern



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China Winter Wheat area, using leading cultivars released from 1949 to 2000. Results showed average annual genetic gain of about 0.31% and 0.74%, respectively. In region 1, yield components did not change though plant height was reduced; in region 2 genetic improvement of grain yield was attributed to the increased thousand kernel weight (0.65%, P < 0.01) and kernel weight/spike (0.87%, P < 0.01). The future challenge of wheat breeding in this region is to continue improving grain yield and disease resistance, and to develop cultivars suitable for the reduced tillage of wheat/rice double cropping. In order to get an overview of the constraints that breeders are facing a survey was conducted covering 19 countries, representing 90% of all wheat grown and produced in less developed countries (Kosina et al). The most signiWcant constraints to wheat production were reported to be heat and water stress, weeds and diseases. Access to mechanization and availability of credit were the socioeconomic constraints most often highlighted. Lack of resources for Weld station operations are an important infrastructural constraint. The most desired outputs from partnerships with international agricultural centers include germplasm development and exchange, assistance in capacity building, and knowledge sharing. The symposium, therefore, simultaneously highlighted reason to be optimistic about improving the impact of wheat breeding through adoption of new technologies, while underscoring the considerable challenges being faced by agricultural researchers due increased demand for wheat as well as environmental and economic constraints. For more information on the plans of the international wheat research community to tackle these issues, readers are referred to the proceedings of symposium (Reynolds et al. 2007b).
Acknowledgments Financial support provided by the Australian Centre for International Agricultural Research (ACIAR) is acknowledged by participants of the International Symposium Challenges to International Wheat Improvement, CIMMYT, Obregon, Mexico, 2024 March 2006, including the authors listed below publishing in this special issue of Euphytica. CIMMYT and CSIRO would also like to acknowledge ACIARs contribution to the project Increasing yield potential in wheat: complementing conventional breeding by application of novel physiological and germplasm strategies (LWR2/1998/014) between 1999 and 2007 that led up to the current forum of international wheat researchers.

Borlaug NE (2007) Sixty-two years of Wghting hunger: personal recollections. Euphytica 157:287297 Brennan JP, Martin PJ (2007) Returns to investment in new breeding technologies. Euphytica 157:337349 Chapman SC, Mathews KL, Trethowan RM, Singh RP (2007) Relationships between height and yield in near-isogenic spring wheats that contrast for major reduced height genes. Euphytica 157:391397 Chatrath R, Mishra B, Ortiz Ferrara G, Singh SK, Joshi AK (2007) Challenges to wheat production in South Asia. Euphytica 157:447456 DePauw RM, Knox RE, Clarke F, Wang H, Fernandez MR, Clarke JM, McCaig TN (2007) Shifting undesirable correlations. Euphytica 157:409415 Duveiller E, Singh RP, Nicol JM (2007) The challenges of maintaining wheat productivity: pests, diseases, and potential epidemics. Euphytica 157:417430 Gupta R, Sayre K (2007) Conservation agriculture in South Asia. J Agric Sci 145:207214 Joshi AK, Mishra B, Chatrath R, Ortiz Ferrara G, Singh RP (2007) Wheat improvement in India: present status, emerging challenges and future prospects. Euphytica 157:431446 Kosina P, Reynolds MP, Dixon J, Joshi AK (2007) Stakeholder perception of wheat production constraints, capacity building needs, and research partnerships in developing countries. Euphytica 157:475483 Ogbonnaya FC, Ye G, Trethowan RM, Dreccer F, Lush D, Shepperd J, van Ginkel M (2007) Yield of synthetic backcross-derived lines in rainfed environments of Australia. Euphytica 157:321336 Ortiz R, Trethowan RM, Ortiz Ferrara G, Iwanaga M, Dodds JH, Crouch JH, Crossa J, Braun HJ (2007) High yield potential, shuttle breeding, genetic diversity, and a new international wheat improvement strategy. Euphytica 157: 365384 Ortiz-Ferrara G, Joshi AK, Chand R, Bhatta MR, Mudwari A, Thapa DB, SuWan MA, Saikia TP, Chatrath R, Witcombe JR, Virk DS, Sharma RC (2007) Partnering with farmers to accelerate adoption of new technologies in South Asia to improve wheat productivity. Euphytica 157: 399407 Reynolds MP, Borlaug NE (2006) Impacts of breeding on international collaborative wheat improvement. J Agric Sci 144:37 Reynolds MP, Hobbs PR, Braun HJ (2007a) Challenges to international wheat improvement. J Agric Sci 145: 223227 Reynolds MP, Poland D, Listman M, Pietragalla J, Braun H (eds) (2007b) Challenges to international wheat improvement. CIMMYT, Mexico City Sharma RC, Ortiz-Ferrara G, Crossa J, Bhatta MR, SuWan MA, Shoran J, Joshi AK, Chand R, Singh G, Ortiz R (2007) Wheat grain yield and stability assessed through regional trials in the Eastern Gangetic Plains of South Asia. Euphytica 157:457464 Singh RP, Huerta-Espino J, Sharma R, Joshi AK, Trethowan RM (2007) High yielding spring bread wheat germplasm


Euphytica (2007) 157:281285 for global irrigated and rainfed production systems. Euphytica 157:351363 Sorrells ME (2007) Application of new knowledge, technologies, and strategies to wheat improvement. Euphytica 157:299306 Trethowan RM, Crossa J (2007) Lessons learnt from forty years of international spring bread wheat trials. Euphytica 157:385390

285 William HM, Trethowan RM, Crosby-Galvan EM (2007) Wheat breeding assisted by markers: CIMMYTs experience. Euphytica 157:307319 Zhou Y, Zhu HZ, Cai SB, He ZH, Zhang XK, Xia XC, Zhang GS (2007) Genetic improvement of grain yield and associated traits in the Southern China winter wheat region: 1949 to 2000. Euphytica 157:465473