Recent progress on molecular breeding of rice in China

Molecular breeding of rice for high yield, superior grain quality, and strong environmental adaptability is crucial for feeding the world’s rapidly growing population. The increasingly cloned quantitative trait loci and genes, genome variations, and haplotype blocks related to agronomically important traits in rice have provided a solid foundation for direct selection and molecular breeding, and a number of genes have been successfully introgressed into mega varieties of rice. Here we summarize China’s great achievements in molecular breeding of rice in the following five traits: high yield, biotic stress resistance, abiotic stress resistance, quality and physiology. Further, the prospect of rice breeding by molecular design is discussed.     

中国水稻遗传育种历程与展望

我国的水稻育种经历了矮化育种、杂种优势利用和绿色超级稻培育3次飞跃,其间伴随矮化育种(第一次绿色革命)、三系杂交稻培育、二系杂交稻培育、亚种间杂种优势利用、理想株型育种和绿色超级稻培育等6个重要历程。育种目标从唯产量是举到高抗、优质和高产并重,育种理念从高产优质逐步提升为“少投入,多产出,保护环境”。水稻功能基因组研究为第二次绿色革命准备了大量的有重要利用价值的基因,水稻育种正迈向设计育种的新时代。基因组选择技术和转基因技术将为培育“少打农药,少施化肥,节水抗旱,优质高产” 绿色超级稻保驾护航。本文对我国水稻遗传育种的发展历程进行了概括,指出了各种育种方法和育种技术的优缺点,系统介绍了水稻细胞质雄性不育和光温敏雄性核不育以及籼粳杂种不育的分子机制的研究进展,综述了水稻株型、穗型、粒形和养分高效利用相关的重要功能基因,阐明了产量与开花期联动的关系,凸显了我国水稻基础研究在国际上的重要地位。特别指出,近年来,我国水稻生产方式发生了或正在发生巨大变革,育种理念也要与时俱进。未来,杂交育种技术要与现代育种技术紧密结合,选育水稻品种不仅要满足市场需求,而且更要具备绿色健康的特点,同时还要适应新耕作制度和新耕作方法。     

Breeding for Yield Potential and Stress Adaptation in Cereals

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, since they include the world's main crops, however, examples from other crops are also included. Topics covered by the review include the conceptual framework for identifying secondary traits associated with yield potential and stress adaptation, and how to measure these secondary traits in practice. The second part of the review deals with the real role of molecular breeding for complex traits from a physiological perspective. This part examines current developments in MAS and quantitative trait loci (QTL) detection as well as plant transformation. Emphasis is placed on the current limitations of these molecular approaches to improving stress adaptation and yield potential. The essay ends by presenting some ideas regarding future avenues for crop breeding given the current and possible future challenges, and on a multidisciplinary approach where physiological knowledge and proper phenotyping play a major role.     

Studies on some genetic parameters of rice (Oryza sativa L.)

Summary In order to utilize the available useful variation in breeding rice to improve yield and quality of grain, the performance of a wide range of rice germ plasm comprising 30 varieties was evaluated in northern India, a major rice growing belt. Plant performance revealed high genetic divergence and phenotypic variability in the crop, with the maximum range of variation being for grain number per panicle and the minimum for grain dimensions. There were also significant differences among varietal means for ten phenotypic traits. Genotypic and phenotypic variance contributed profoundly to the variance of the phenotypic traits studied, but, since genetic variability in the traits related to yield was considerable, there is scope for further improvement in yielding ability. Grain number per panicle, number of effective tillers per plant and culm length exhibit high heritability, and genotypic coefficient of variation and therefore a high genetic advance. Thus, selection for these traits would be effective in crop improvement. Moreover, grain number per panicle shows a significant positive correlation with yield, and this trait could profitably form a reliable index for the yielding capacity of this crop.     

Inducing drought tolerance in plants: Recent advances

Undoubtedly, drought is one of the prime abiotic stresses in the world. Crop yield losses due to drought stress are considerable. Although a variety of approaches have been used to alleviate the problem of drought, plant breeding, either conventional breeding or genetic engineering, seems to be an efficient and economic means of tailoring crops to enable them to grow successfully in drought-prone environments. During the last century, although plant breeders have made ample progress through conventional breeding in developing drought tolerant lines/cultivars of some selected crops, the approach is, in fact, highly time-consuming and labor- and cost-intensive. Alternatively, marker-assisted breeding (MAB) is a more efficient approach, which identifies the usefulness of thousands of genomic regions of a crop under stress conditions, which was, in reality, previously not possible. Quantitative trait loci (QTL) for drought tolerance have been identified for a variety of traits in different crops. With the development of comprehensive molecular linkage maps, marker-assisted selection procedures have led to pyramiding desirable traits to achieve improvements in crop drought tolerance. However, the accuracy and preciseness in QTL identification are problematic. Furthermore, significant genetic × environment interaction, large number of genes encoding yield, and use of wrong mapping populations, have all harmed programs involved in mapping of QTL for high growth and yield under water limited conditions. Under such circumstances, a transgenic approach to the problem seems more convincing and practicable, and it is being pursued vigorously to improve qualitative and quantitative traits including tolerance to biotic and abiotic stresses in different crops. Rapid advance in knowledge on genomics and proteomics will certainly be beneficial to fine-tune the molecular breeding and transformation approaches so as to achieve a significant progress in crop improvement in future. Knowledge of gene regulation and signal transduction to generate drought tolerant crop cultivars/lines has been discussed in the present review. In addition, the advantages and disadvantages as well as future prospects of each breeding approach have also been discussed.     

Molecular approaches for genetic improvement of seed quality and characterization of genetic diversity in soybean: a critical review

Soybean is an economically important leguminous crop. Genetic improvements of soybeans have focused on enhancement of seed and oil yield, development of varieties suited to different cropping systems, and breeding resistant/tolerant varieties for various biotic and abiotic stresses. Plant breeders have used conventional breeding techniques for the improvement of these traits in soybean. The conventional breeding process can be greatly accelerated through the application of molecular and genomic approaches. Molecular markers have proved to be a new tool in soybean breeding by enhancing selection efficiency in a rapid and time-bound manner. An overview of molecular approaches for the genetic improvement of soybean seed quality parameters, considering recent applications of marker-assisted selection and ‘omics’ research, is provided in this article.     

Transmission of important chromosomal regions under selection revealed in rice pedigree breeding programs

Genetic analysis across a whole plant genome based on pedigree information offers considerable potential for enhancing genetic gain from plant breeding programs through quantitative trait loci (QTL) mapping and marker-assisted selection. Here, we report its application for graphically genotyping varieties used in Chinese japonica rice (Oryza sativa L.) pedigree breeding programs. We identified 34 important chromosomal regions from the founder parent that are under selection in the breeding programs, and by comparing donor genomic regions that are under selection with QTL locations of agronomic traits, we found that QTL clustered in important genomic regions, in accordance with association analyses of natural populations and other previous studies. The convergence of genomic regions under selection with QTL locations suggests that donor genomic regions harboring key genes/QTL for important agronomic traits have been selected by plant breeders since the 1950s from the founder rice plants. The results provide better understanding of the effects of selection in breeding programs on the traits of rice cultivars. They also provide potentially valuable information for enhancing rice breeding programs through screening candidate parents for targeted molecular markers, improving crop yield potential and identifying suitable genetic material for use in future breeding programs.     

Yield-related QTLs and Their Applications in Rice Genetic Improvement

Grain yield is one of the most important indexes in rice breeding,which is governed by quantitative trait loci (QTLs).Different map-ping populations have been used to explore the QTLs controlling yield related traits.Primary populations such as F 2 and recombi-nant inbred line populations have been widely used to discover QTLs in rice genome-wide,with hundreds of yield-related QTLs detected.Advanced populations such as near isogenic lines (NILs) are efficient to further fine-map and clone target QTLs.NILs for primarily identified QTLs have been proposed and confirmed to be the ideal population for map-based cloning.To date,20 QTLs directly affecting rice grain yield and its components have been cloned with NIL-F 2 populations,and 14 new grain yield QTLs have been validated in the NILs.The molecular mechanisms of a continuously increasing number of genes are being unveiled,which aids in the understanding of the formation of grain yield.Favorable alleles for rice breeding have been ’mined’ from natural cultivars and wild rice by association analysis of known functional genes with target trait performance.Reasonable combination of favorable alleles has the potential to increase grain yield via use of functional marker assisted selection.     

赤霉素生物合成与信号传递对植物株高的调控

植物株高是影响作物产量和品质的重要农艺性状。赤霉素(Gibberellins,GAs)是调控植物株高的重要激素,GA相关株高基因的克隆与分子机制研究对于合理调控作物生长发育和农业生产具有极其重要的利用价值,在水稻、小麦等粮食作物育种中得到了广泛应用。为了促进GA在果树、花卉等园艺作物育种中的有效利用,文中在分子生物学水平上介绍GA生物合成和GA信号传递途径对植物株高的调控。     

Congruence between selection on breeding values and farmers' selection criteria in sheep breeding under conventional nucleus breeding schemes

Designing breeding schemes suitable for smallholder livestock production systems in developing regions has hitherto been a challenge. The suggested schemes either do not address farmers' breeding goals (centralized station-based nucleus schemes) or yield slow genetic progress (village-based schemes). A new breeding scheme that integrates the merits of previously suggested schemes has been designed for Menz sheep improvement in Ethiopia. It involves selection based on breeding values in nucleus flocks to produce elite rams, a one-time only provision of improved rams to villagers and a follow-up village-based selection to sustain genetic progress in village flocks. Here, we assessed whether conventional selection of breeding rams based on breeding values for production traits, which is the practice in station-based nucleus flocks, meets farmers' breeding objectives. We also elicited determinants of farmers' ram choice. Low but significant correlations were found between rankings of rams based on farmers' selection criteria, estimated breeding values (EBV) and body weight (BW). Appearance traits (such as color and horn) and meat production traits (BW and linear size traits) significantly determined farmers' breeding ram choice. The results imply that conventional selection criteria based solely on EBV for production traits do not address farmers' trait preferences fully, but only partially. Thus, a two-stage selection procedure involving selection on breeding values in nucleus centers followed by farmers' selection among top- ranking candidate rams is recommended. This approach accommodates farmers' preferences and speeds up genetic progress in village-based selection. The Menz sheep scheme could be applied elsewhere with similar situations to transform conventional station-based nucleus breeding activities into participatory breeding programs.     

Genetic basis of yield as viewed from a crop physiologist's perspective

The final yield of a crop is the product of growth during the growing season and a number of developmental processes occurring throughout the life cycle of a crop, with most genes influencing the final outcome to a degree. However, recent advances in molecular biology have developed the potential to identify and map many genes or QTLs related to various important traits, including yield, plant adaptation and tolerance to stresses. Significant G×E interactions for yield have been identified, as have interactions associated with QTLs for yield. However, there is little evidence available to confirm that a QTL for yield from a parental line in one mapping population may improve yield when transferred into an adapted, high‐yielding line of another population. In order to narrow the apparent gap between the genotype and the phenotype with regard to yield, it is important to identify key traits related to yield and then attempt to identify and locate the genes controlling them. The partitioning of the developmental time to anthesis into different phases: from sowing to the onset of stem elongation and from then to anthesis, as a relatively simple physiological attribute putatively related to yield, is discussed. If the relationship holds in a wider range of conditions and the genetic factors responsible are located then the genetic basis of yield should be identified. There has also been significant progress in crop simulation modelling. Using knowledge of crop physiology and empirical relationships these models can simulate the performance of crops, including the G×E interactions. Such models require information regarding the genetic basis of yield, which are included in the form of genetic coefficients. Essentially models are constructed as decision‐making tools for management but may be of use in detecting prospective traits for selection within a breeding programme. Problems associated with this approach are discussed. This review discusses the need to use crop physiology approaches to analyse components of yield in order to reliably identify the genetic basis of yield.     

Advances to improve the eating and cooking qualities of rice by marker-assisted breeding

The eating and cooking qualities of rice are heavily emphasized in breeding programs because they determine market values and they are the appealing attributes sought by consumers. Conventional breeding has developed traditional varieties with improved eating and cooking qualities. Recently, intensive genetic studies have pinpointed the genes that control eating and cooking quality traits. Advances in genetic studies have developed molecular techniques, thereby allowing marker-assisted breeding (MAB) for improved eating and cooking qualities in rice. MAB has gained the attention of rice breeders for the advantages it can offer that conventional breeding cannot. There have been successful cases of using MAB to improve the eating and cooking qualities in rice over the years. Nevertheless, MAB should be applied cautiously given the intensive effort needed for genotyping. Perspectives from conventional breeding to marker-assisted breeding will be discussed in this review for the advancement of the eating and cooking qualities of fragrance, amylose content (AC), gel consistency (GC) and gelatinization temperature (GT) in rice. These four parameters are associated with eating and cooking qualities in rice. The genetic basis of these four parameters is also included in this review. MAB is another approach to rice variety improvement and development in addition to being an alternative to genetic engineering. The MAB approach shortens the varietal development time, and is therefore able to deliver improved rice varieties to farmers within a shorter period of time.     

What it will take to Feed 5.0 Billion Rice consumers in 2030

Major advances have occurred in rice production due to adoption of green revolution technology. Between 1966 and 2000, the population of densely populated low income countries grew by 90% but rice production increased by 130% from 257 million tons in 1966 to 600 million tons in 2000. However, the population of rice consuming countries continues to grow and it is estimated that we will have to produce 40 more rice in 2030. This increased demand will have to be met from less land, with less water, less labor and fewer chemicals. To meet the challenge of producing more rice from suitable lands we need rice varieties with higher yield potential and greater yield stability. Various strategies for increasing the rice yield potential being employed include: (1) conventional hybridization and selection procedures, (2) ideotype breeding, (3) hybrid breeding, (4) wide hybridization and (5) genetic engineering. Various conventional and biotechnology approach are being employed to develop durable resistance to diseases and insect and for tolerance to abiotic stresses. The availability of the rice genome sequence will now permit identification of the function of each of 60,000 rice genes through functional genomics. Once the function of a gene is identified, it will be possible to develop new rice varieties by introduction of the gene through traditional breeding in combination with marker aided selection or direct engineering of genes into rice varieties.     

A whole‐genome SNP array (RICE6K) for genomic breeding in rice

The advances in genotyping technology provide an opportunity to use genomic tools in crop breeding. As compared to field selections performed in conventional breeding programmes, genomics‐based genotype screen can potentially reduce number of breeding cycles and more precisely integrate target genes for particular traits into an ideal genetic background. We developed a whole‐genome single nucleotide polymorphism (SNP) array, RICE6K, based on Infinium technology, using representative SNPs selected from more than four million SNPs identified from resequencing data of more than 500 rice landraces. RICE6K contains 5102 SNP and insertion–deletion (InDel) markers, about 4500 of which were of high quality in the tested rice lines producing highly repeatable results. Forty‐five functional markers that are located inside 28 characterized genes of important traits can be detected using RICE6K. The SNP markers are evenly distributed on the 12 chromosomes of rice with the average density of 12 SNPs per 1 Mb and can provide information for polymorphisms between indica and japonica subspecies as well as varieties within indica and japonica groups. Application tests of RICE6K showed that the array is suitable for rice germplasm fingerprinting, genotyping bulked segregating pools, seed authenticity check and genetic background selection. These results suggest that RICE6K provides an efficient and reliable genotyping tool for rice genomic breeding.     

Field performance of STG06L-35-061, a new genetic resource developed from crosses between weed-suppressive indica rice and commercial southern U.S. long-grains

Aims

Weed control in rice is challenging, particularly in light of increased resistance to herbicides in weed populations including Echinochloa crus-galli (L.) Beauv. Indica rice cultivars can produce high yields and suppress barnyardgrass, but have not been commercially acceptable in the U.S. due to inferior agronomic traits and grain quality. Our objectives were to combine high yield and weed-suppressive characteristics from indica cultivars with commercially acceptable grain quality and plant types from long-grain cultivars grown in the southern U.S.

Methods

Crosses between indica and commercial tropical japonica (cv. Katy, and cv. Drew) rice were evaluated for weed suppression and agronomic traits in a breeding program.

Results

In some tests, the selection STG06L-35-061 was nearly as weed suppressive as PI 312777, the suppressive parent, and more suppressive than its tropical japonica parents. Its main crop yield is commercially acceptable, and intermediate between PI 312777 and Katy. Its milling quality and cooking quality are similar to long-grain commercial cultivars, and it has resistance to rice blast disease. Marker analyses identified introgressions from the indica parents on chromosomes 1 and 3 of STG06L-35-061 that require further analysis as possible sources of weed suppressive traits.

Conclusions

STG06L-35-061 might be suitable for organic rice or reduced input conventional systems.     

作物基因聚合分子育种

基因聚合分子育种与常规育种技术相结合已成为今后作物育种的主流方向。基因聚合分子育种主要包括遗传转化基因聚合分子育种和分子标记筛选基因聚合分子育种。本文简要综述了近年来作物基因聚合分子育种的研究进展,分析了遗传转化基因聚合分子育种以及分子标记基因聚合分子育种技术的研究方法及基因聚合分子育种存在的问题。     

Breeding implications of drought stress under future climate for upland rice in Brazil

Rice is the most important food crop in the developing world. For rice production systems to address the challenges of increasing demand and climate change, potential and on‐farm yield increases must be increased. Breeding is one of the main strategies toward such aim. Here, we hypothesize that climatic and atmospheric changes for the upland rice growing period in central Brazil are likely to alter environment groupings and drought stress patterns by 2050, leading to changing breeding targets during the 21st century. As a result of changes in drought stress frequency and intensity, we found reductions in productivity in the range of 200–600 kg/ha (up to 20%) and reductions in yield stability throughout virtually the entire upland rice growing area (except for the southeast). In the face of these changes, our crop simulation analysis suggests that the current strategy of the breeding program, which aims at achieving wide adaptation, should be adjusted. Based on the results for current and future climates, a weighted selection strategy for the three environmental groups that characterize the region is suggested. For the highly favorable environment (HFE, 36%–41% growing area, depending on RCP), selection should be done under both stress‐free and terminal stress conditions; for the favorable environment (FE, 27%–40%), selection should aim at testing under reproductive and terminal stress, and for the least favorable environment (LFE, 23%–27%), selection should be conducted for response to reproductive stress only and for the joint occurrence of reproductive and terminal stress. Even though there are differences in timing, it is noteworthy that stress levels are similar across environments, with 40%–60% of crop water demand unsatisfied. Efficient crop improvement targeted toward adaptive traits for drought tolerance will enhance upland rice crop system resilience under climate change.     

Marker assisted selection in crop plants

Genetic mapping of major genes and quantitative traits loci (QTLs) for many important agricultural traits is increasing the integration of biotechnology with the conventional breeding process. Exploitation of the information derived from the map position of traits with agronomical importance and of the linked molecular markers, can be achieved through marker assisted selection (MAS) of the traits during the breeding process. However, empirical applications of this procedure have shown that the success of MAS depends upon several factors, including the genetic base of the trait, the degree of the association between the molecular marker and the target gene, the number of individuals that can be analyzed and the genetic background in which the target gene has to be transferred. MAS for simply inherited traits is gaining increasing importance in breeding programs, allowing an acceleration of the breeding process. Traits related to disease resistance to pathogens and to the quality of some crop products are offering some important examples of a possible routinary application of MAS. For more complex traits, like yield and abiotic stress tolerance, a number of constraints have determined severe limitations on an efficient utilization of MAS in plant breeding, even if there are a few successful applications in improving quantitative traits. Recent advances in genotyping technologies together with comparative and functional genomic approaches are providing useful tools for the selection of genotypes with superior agronomical performancies.