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The development of germplasm resources and advances in breeding methods have led to steady increases in yield and quality of rice (Oryza sativa L.). Three milestones in the recent history of rice breeding have contributed to these increases: dwarf rice breeding, hybrid rice breeding, and super rice breeding. On the 50th anniversary of the success of three-line hybrid rice, we highlight important scientific discoveries in rice breeding that were made by Chinese scientists and summarize the broader history of the field. We discuss the strategies that could be used in the future to optimize rice breeding further in the hope that China will continue to play a leading role in international rice breeding.

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Enhancing photosynthetic capacity is widely accepted as critical to advancing crop yield. Therefore, identifying photosynthetic parameters positively related to biomass accumulation in elite cultivars is the major focus of current rice research. In this work, we assessed leaf photosynthetic performance, canopy photosynthesis, and yield attributes of super hybrid rice cultivars Y-liangyou 3218 (YLY3218) and Y-liangyou 5867 (YLY5867) at tillering stage and flowering stage, using inbred super rice cultivars Zhendao11(ZD11) and Nanjing 9108 (NJ9108) as control. A diurnal canopy photosynthesis model was applied to estimate the influence of key environmental factors, canopy attributes, and canopy nitrogen status on daily aboveground biomass increment (AMDAY). Results showed that primarily the light-saturated photosynthetic rate at tillering stage contributed to the advancing yield and biomass of super hybrid rice in comparison to inbred super rice, and the light-saturated photosynthetic rate between them was similar at flowering stage. At tillering stage, the higher CO2 diffusion capacity, together with higher biochemical capacity (i.e., maximum carboxylation rate of Rubisco, maximum electron transport rate (J max), and triose phosphate utilization rate) favored leaf photosynthesis of super hybrid rice. Similarly, AMDAY in super hybrid rice was higher than inbred super rice at tillering stage, and comparable at flowering stage partially due to increased canopy nitrogen concentration (SLNave) of inbred super rice. At tillering stage, model simulation revealed that replacement of J max and g m in inbred super rice by super hybrid rice always had a positive effect on AMDAY, and the averaged AMDAY increment was 5.7% and 3.4%, respectively. Simultaneously, the 20% enhancement of total canopy nitrogen concentration through the improvement of SLNave (TNC-SLNave) resulted in the highest AMDAY across cultivars, with an average increase of 11.2%. In conclusion, the advancing yield performance of YLY3218 and YLY5867 was due to the higher J max and g m at tillering stage, and TCN-SLNave is a promising target for future super rice breeding programs.

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Rice (Oryza sativa) is an important staple food crop worldwide, especially in east and southeast Asia. About one-third of rice cultivated area is under saline soil, either natural saline soils or irrigation with brackish water. Salinity stress is among the devastating abiotic stresses that not only affect rice growth and crop productivity but also limit its cultivation area globally. Plants adopt multiple tolerance mechanisms at the morphological, physiological, and biochemical levels to tackle salinity stress. To identify these tolerance mechanisms, this study was carried out under both a controlled glass house as well as natural saline field conditions using 22 green super rice (GSR) lines along with two local varieties ("IRRI 6 and Kissan Basmati"). Several morpho-physiological and biochemical parameters along with stress-responsive genes were used as evaluation criteria under normal and salinity stress conditions. Correlation and Principal Component Analysis (PCA) suggested that shoot-related parameters and the salt susceptible index (SSI) can be used for the identification of salt-tolerant genotypes. Based on Agglomerative Hierarchical Cluster (AHC) analysis, two saline-tolerant ("S19 and S20") and saline-susceptible ("S3 and S24") lines were selected for further molecular evaluation. Quantitative RT-PCR was performed, and results showed that expression of 1-5-phosphoribosyl -5-5-phosphoribosyl amino methylidene amino imidazole-4-carboxamide isomerase, DNA repair protein recA, and peptide transporter PTR2 related genes were upregulated in salt-tolerant genotypes, suggesting their potential role in salinity tolerance. However, additional validation using reverse genetics approaches will further confirm their specific role in salt tolerance. Identified saline-tolerant lines in this study will be useful genetic resources for future salinity breeding programs.

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It is well documented that yield superiority of super hybrid rice is linked with its improved photosynthetic capacity and/or efficiency. In natural environments, the amounts of CO2 assimilated by plants was also impacted by the rapidity of leaf photosynthesis response to fluctuations of light. However, it remains unknow whether the high yield of super hybrid rice was associated with photosynthetic traits under dynamic state. Here, photosynthetic traits under steady-and dynamic state in two super hybrid rice varieties (Ylinagyou 3218 and Yliangyou 5867) with high yield and two inbred super rice varieties (Zhendao 11 and Nanjing 9108) with lower yield. Under steady state, the net photosynthetic rate (A*) in super hybrid rice was 25.3% larger compared with inbred super rice. During photosynthetic induction, there was no obvious association of the rapidity of net photosynthesis rate (A) to sunflecks with rice subpopulations. Stomatal conductance (gs) of super hybrid rice increased slower than that of inbred super rice. The cumulative CO2 fixation (CCF) during photosynthetic induction was 25.2% larger in super hybrid rice than that in inbred super rice. The primary limitation during induction was biochemical limitation rather than stomatal limitation. There was a significantly positive relationship between A* and CCF, while A* was not related with the induction response rate of A. Overall, A* and CCF in super hybrid rice have been improved together, which contributed to its yield superiority, whereas its yield potential still can be improved by increasing induction rate of A under fluctuations of irradiance.

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Producing sufficient food with finite resources to feed the growing global population while having a smaller impact on the environment has always been a great challenge. Here, we review the concept and practices of Green Super Rice (GSR) that have led to a paradigm shift in goals for crop genetic improvement and models of food production for promoting sustainable agriculture. The momentous achievements and global deliveries of GSR have been fueled by the integration of abundant genetic resources, functional gene discoveries, and innovative breeding techniques with precise gene and whole-genome selection and efficient agronomic management to promote resource-saving, environmentally friendly crop production systems. We also provide perspectives on new horizons in genomic breeding technologies geared toward delivering green and nutritious crop varieties to further enhance the development of green agriculture and better nourish the world population.

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To investigate the differential responses of super rice grain filling to low filling stage temperature (LT) and the regulative effect of nitrogen panicle fertilizer (NPF), physiological and molecular experiments were conducted with two super rice varieties (Nanjing 7th: N7 and Nanjing 9108th: N9108) on two different filling stage temperature treatments implemented by applying two sowing dates [Normal filling stage temperature (CK): Sowed on May 30, Tmean = 24.7°C and low filling stage temperature (LT): Sowed on July 1, Tmean = 20.3°C], and two NPF levels (0 and 150 kg N ha-1). In this study, LT, NPF, and simultaneous LT and NPF treatments suppressed the grain filling in all varieties with different levels. Under LT or NPF treatments, the reduction of grain weight, seed setting rate, and filling rate were closely associated with suppressed starch biosynthesis rate in inferior seeds, suggesting that reduced starch biosynthesis rate, expression, and activities of enzymes encoded by related genes, Floury endosperm-4 (FLO4), Starch branching enzyme-I (SBE1), and Starch phosphorylase-L (PHO-l), were responsible for the grain filling reduction. Under LT or NPF treatments, significantly higher grain filling rates and lower variance were found in N9108 compared to that in N7, which were closely related to their higher starch biosynthesis ability, related gene expression, and enzymes activities. One of the probable explanations of the grain filling difference was the variation in the relative amount of key regulative hormones, Abscisic acid (ABA) and 1-aminocyclopropane-1-carboxylic acid (ACC). These results raise a possibility that super rice with higher sink activities has superior adaptability to LT and NPF due to their higher sink activities.

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Great progress has been achieved in developing hybrid "super rice" varieties in China. Understanding morphological root traits in super rice and the mechanisms of nitrogen acquisition by the root system are of fundamental importance to developing proper fertilisation and nutrient management practices in their production. The present study was designed to study morphological and physiological traits in hybrid super rice roots that are associated with nitrogen use efficiency (NUE). Two hybrid super rice varieties (Yongyou12, YY; Jiayou 6, JY) and one common variety (Xiushui 134, XS) with differing NUE were cultivated hydroponically, and morphological and physiological traits of seedling roots in response to varying nitrogen conditions were investigated. Our results show that the hybrid cultivars YY and JY exhibit larger root systems, arising from a maximisation of root tips and from longer roots without changes in root diameter. The cross-sectional proportion of aerenchyma was significantly higher in super rice roots. The larger root system of super hybrid rice contributed to higher N accumulation and resulted in higher N uptake efficiency. 15N (15NH4+) labeling results show that YY and JY had an enhanced capacity for ammonium (NH4+) uptake. Moreover, YY and JY were more tolerant to high NH4+ and showed reduced futile NH4+ efflux. NH4+ efflux in the root elongation zone, measured by Non-invasive Micro-test Technology, was significantly lower than in XS. Taken together, our results suggest that a longer root, a larger number of tips, a better developed aerenchyma, a higher capacity for N uptake, and reduced NH4+ efflux from roots are associated with higher NUE and growth performance in hybrid super rice.

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Food security is confronted by major threats from crop yield stagnation and global climate change. The benefits of phenotypic plasticity across environments for given crop genotypes are thought to be imperative for high-yielding cropping systems. Given that 3D modeling is increasingly recognized for dissecting crop phenotypic plasticity, it requires an assessment of the potential benefits of architectural adaptation of super rice to different agronomic practices. In this study, we focused on a comprehensive evaluation of the phenotypic plasticity of super rice on the aspects of 3D architectural "reoptimization," photosynthetic productivity, nitrogen economy, and grain yield. A super rice phenotype in superhigh-yielding practice (SH) displays a "reoptimized" morphogenesis in the leaf vertical dispersion and orientation in comparison to that in Farmer's practice (FP). Specifically, a super rice phenotype in SH is provided with a high cumulative rate and peaks of leaf area, increasing the distribution of high leaf inclination angles in comparison to that in FP, particularly in the upper parts of the canopy. These "reoptimizations" sustained profits in light environment within a canopy, leaf area duration, photosynthetic light harvest, and light utilization efficiency and were coordinated with improving nitrogen uptake and assimilation. The current literature indicates that the agronomic plasticity of super rice in architectural "reoptimization" is a promising perspective for high yield formation. Our results suggest that more emphasis should be placed upon agronomic adaptation strategies for super rice across diverse genotypes and environments to further improve crop establishment and photosynthetic productivity.

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Drought and low nitrogen are the most common abiotic stresses limiting rice productivity in the rainfed rice areas of Asia and Africa. Development and adoption of green super rice (GSR) varieties with greatly improved drought tolerance (DT) and low nitrogen tolerance (LNT) are the most efficient way to resolve this problem. In this study, using three sets of trait-specific introgression lines (ILs) in a Xian (indica) variety Huanghuazhan (HHZ) background, we identified nine DT-QTL and seven LNT-QTL by a segregation distortion approach and a genome-wide association study, respectively. Based on performances of DT and LNT and genotypes at the detected QTL, two ILs M79 and M387 with DT and LNT were selected for cross-making to validate the identified QTL and to develop DT and LNT rice lines by pyramiding two DT-QTL (qDT3.9 and qDT6.3) and two LNT-QTL (qGY1 and qSF8). Using four pairs of kompetitive allele specific PCR (KASP) SNP markers, we selected 66 F2 individuals with different combinations of the target DT- and LNT-QTL favorable alleles and they showed expected improvement in DT and/or LNT, which were further validated by the significant improvement in DT and/or LNT of their F3 progeny testing. Based on evaluation of pyramiding lines in F3 lines under drought, low nitrogen (LN) and normal conditions, four promising pyramiding lines having different QTL favorable alleles were selected, which showed significantly improved tolerances to drought and/or LN than HHZ and their IL parents. Our results demonstrated that trait-specific ILs could effectively connect QTL mapping and QTL pyramiding breeding, and designed QTL pyramiding (DQP) using ILs could be more effective in molecular rice breeding for complex quantitative traits.

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Rice (Oryza sativa) is a major staple food crop for more than 3.5 billion people worldwide. Understanding the regulatory mechanisms of complex agronomic traits in rice is critical for global food security. Rice is also a model plant for genomics research of monocotyledons. Thanks to the rapid development of functional genomic technologies, over 2000 genes controlling important agronomic traits have been cloned, and their molecular biological mechanisms have also been partially characterized. Here, we briefly review the advances in rice functional genomics research during the past 10 years, including a summary of functional genomics platforms, genes and molecular regulatory networks that regulate important agronomic traits, and newly developed tools for gene identification. These achievements made in functional genomics research will greatly facilitate the development of green super rice. We also discuss future challenges and prospects of rice functional genomics research.

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Breeding of multi-stress tolerant rice varieties with higher grain yields is the best option to enhance the rice productivity of abiotic stresses prone areas. It also poses the greatest challenge to plant breeders to breed rice varieties for such stress prone conditions. Here, we carried out a designed QTL pyramiding experiment to develop high yielding "Green Super Rice" varieties with significantly improved tolerance to salt stress and grain yield. Using the F4 population derived from a cross between two selected introgression lines, we were able to develop six mostly homozygous promising high yielding lines with significantly improved salt tolerance and grain yield under optimal and/or saline conditions in 3 years. Simultaneous mapping using the same breeding population and tunable genotyping-by-sequencing technology, we identified three QTL affecting salt injury score and leaf chlorophyll content. By analyzing 32M SNP data of the grandparents and graphical genotypes of the parents, we discovered 87 positional candidate genes for salt tolerant QTL. According to their functional annotation, we inferred the most likely candidate genes. We demonstrated that designed QTL pyramiding is a powerful strategy for simultaneous improvement and genetic dissection of complex traits in rice.

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Selecting rice varieties with a high nitrogen (N) use efficiency (NUE) is the best approach to reduce N fertilizer application in rice production and is one of the objectives of the Green Super Rice (GSR) Project in China. However, the performance of elite candidate GSR varieties under low N supply remains unclear. In the present study, differences in the grain yield and NUE of 13 and 14 candidate varieties with two controls were determined at a N rate of 100 kg ha(-1) in field experiments in 2014 and 2015, respectively. The grain yield for all of the rice varieties ranged from 8.67 to 11.09 t ha(-1), except for a japonica rice variety YG29, which had a grain yield of 6.42 t ha(-1). HY549 and YY4949 produced the highest grain yield, reflecting a higher biomass production and harvest index in 2014 and 2015, respectively. Total N uptake at maturity (TNPM) ranged from 144 to 210 kg ha(-1), while the nitrogen use efficiency for grain production (NUEg) ranged from 35.2 to 62.0 kg kg(-1). Both TNPM and NUEg showed a significant quadratic correlation with grain yield, indicating that it is possible to obtain high grain yield and NUEg with the reduction of TNPM. The correlation between N-related parameters and yield-related traits suggests that promoting pre-heading growth could increase TNPM, while high biomass accumulation during the grain filling period and large panicles are important for a higher NUEg. In addition, there were significant and negative correlations between the NUEg and N concentrations in leaf, stem, and grain tissues at maturity. Further improvements in NUEg require a reduction in the stem N concentration but not the leaf N concentration. The daily grain yield was the only parameter that significantly and positively correlated with both TNPMand NUEg. This study determined variations in the grain yield and NUE of elite candidate GSR rice varieties and provided plant traits that could be used as selection criteria in breeding N-efficient rice varieties.