Changes in Enzyme Activities Involved in Starch Synthesis and Hormone Concentrations in Superior and Inferior Spikelets and Their Association with Grain Filling of Super Rice

The changes in activities of key enzymes involved in sucrose-to-starch conversion and concentrations of hormones in superior and inferior spikelets of super rice were investigated and their association with grain filling was analyzed.Four super rice cultivars,Liangyoupeijiu,IIyou 084,Huaidao 9 and Wujing 15,and two high-yielding and elite check cultivars,Shanyou 63 and Yangfujing 8,were used.The activities of sucrose synthase (SuSase),adenosine diphosphoglucose pyrophosphorylase (AGPase),starch synthase (StSase) and starch branching enzyme (SBE),and the concentrations of zeatin + zeatin riboside (Z + ZR),indole-3-acetic acid (IAA) and abscisic acid (ABA) in superior and inferior spikelets were determined during the grain filling period and their relationships with grain filling rate were analyzed.Maximum grain filling rate,the time reaching the maximum grain-filling rate,mean grain filling rate and brown rice weight for superior spikelets showed a slight difference between the super and check rice cultivars,but were significantly lower in the super rice than in the check rice for inferior spikelets.Changes of enzyme activities and hormone concentrations in grains exhibited single peak curves during the grain filling period.The peak values and the mean activities of SuSase,AGPase,StSase and SBE were lower in inferior spikelets than in superior ones,as well as the peak values and the mean concentrations of Z + ZR and IAA.However,the peak value and the mean concentration of ABA were significantly higher in inferior spikelets than in superior ones and greater in the super rice than in the check rice.The grain filling rate was positively and significantly correlated with the activities of SuSase,AGPase and StSase and the concentrations of Z + ZR and IAA.The results suggested that the low activities of SuSase,AGPase and StSase and the low concentrations of Z + ZR and IAA might be important physiological reasons for the slow grain filling rate and light grain weight of inferior spikelets in super rice.     

Difference in Grain Yield and Quality among Tillers in Rice Genotypes Differing in Tillering Capacity

A japonica variety and its iso-allelic mutant with high tillering ability were used to investigate the differences in grain yield and quality among different tillers. There was a distinct difference in panicle weight among tillers during grain filling for both genotypes, with Xiushui 11 having a greater increase rate in panicle weight, and being earlier in reaching the maximum panicle weight than the mutant. There was a great significant difference between the two genotypes in grain yield and its components, with Xiushui 11 having higher grain yield, more grains per panicle, higher filled grain percentage and grain weight than the mutant for each type of tiller. Moreover, a significant difference was found in grain yield and yield components among different tillers for both genotypes, with grain number per panicle showing the greatest variation over tillers among all yield components. Compared with Xiushui 11, the mutant had higher brown rice rate and greater ratio of length to width of brown rice, and lower chalky rice rate, amylose content, and protein content of rice. Furthermore, there was a significant difference in grain quality among tillers within a plant for both genotypes, with later initiated tillers being lower in chalky rice rate, amylose and protein contents than early initiated ones. The variation of most quality parameters among tillers within a plant was markedly larger for the mutant than for Xiushui 11.     

Changes in Grain Yield of Rice and Emission of Greenhouse Gases from Paddy Fields after Application of Organic Fertilizers Made from Maize Straw

A field experiment was conducted at the farm of Yangzhou University, Yangzhou, China, to study the effects of organic fertilizers made from maize straw on rice grain yield and the emission of greenhouse gases. Four organic fertilizer treatments were as follows： maize straw （MS）, compost made from maize straw （MC）, methane-generating maize residue （MR）, and black carbon made from maize straw （BC）. These organic fertilizers were applied separately to paddy fields before rice transplanting. No organic fertilizer was applied to the control （CK）. The effects of each organic fertilizer on rice grain yield and emission of greenhouse gases were investigated under two conditions, namely, no nitrogen （N） application （ON） and site-specific N management （SSNM）. Rice grain yields were significantly higher in the MS, MC and MR treatments than those in CK under either ON or SSNM. The MS treatment resulted in the highest grain yield and agronomic N use efficiency. However, no significant difference was observed for these parameters between the BC treatment and CK. The changes in the emissions of methane （CH4） carbon dioxide （CO2）, or nitrous oxide （N20） from the fields were similar among all organic fertilizer treatments during the entire rice growing season. The application of each organic fertilizer significantly increased the emission of each greenhouse gas （except N20 emission in the BC treatment） and global warming potential （GWP）. Emissions of all the greenhouse gases and GWP increased under the same organic fertilizer treatment in the presence of N fertilizer, whereas GWP per unit grain yield decreased. The results indicate that the application of organic fertilizer （MS, MC or MR） could increase grain yield, but also could enhance the emissions of greenhouse gases from paddy fields. High grain yield and environmental efficiency could be achieved by applying SSNM with MR.     

Exploitation of genetic variation caused by space treatment in rice

Seeds of early indica rice varieties 6188, D6049, of late indiea rice varieties Minghui 63, Xiangge, Waiqi, 70321, and late japonica rice variety Beik 15 were carried by the retrieved satellite "921006" in order to investigate the comprehensive effects of the space factors. Treated seeds and untreated seeds (CK) were grown in the field. The results showed that although compared with the CK (which was very uniform) small differences on population uniformity, plant height, growth duration and seed setting rate were observed, no segregation was found in SP_1. Strong segregation occurred in SP_2 and SP_3 generations. Significant difference on the grain yield of Single plant was observed between SP_2 and CK, while the yield between     

Practice and Thought on Developing Hybrid Rice for Super High Yield by Exploiting Inter-subspecific Heterosis

Since the breakthrough of grain yield owing to the development of dwarf rice and three-line system hybrid rice, rice breeding for high yield hardly had showed significant progress in the next successive two decades. It was considered that utilizing heterosis between subspecific varieties (Oryza sativa L.) would be an effective approach to increase yield further. During 1987-1993,an indica-japonica hybrid Yayou 2 yielded as high as 10.5 t/ha; however, it failed to be commercialized because of seed purity problem due to non-uniform emasculation by chemical agent in seed production, and sensitivity of seed setting in F1 plants to environmental conditions. In the past decade, two inter-subspeific hybrids, Liangyoupeijiu (Peiai 64S/9311, javanica/indica) and Liangyou E32 (Peiai 64S/E32, javanica/japonica); both of them exhibited grain yield higher than 10.5 t/ha, and were widely judged as the pioneers of super hybrid rice. Liangyoupeijiu has been successfully popularized over 4 million hectare in wide climatic areas, while Liangyou E32 made a yield record and offered a model of plant ideotype for super hybrid rice. It was considered that in combination with plant ideotype, active physiological functions, and wide-range adaptability to ecological conditions, exploitation of indica-japonica heterosis would be the key approach for super hybrid rice breeding.     

Practices and Prospects of Super Hybrid Rice Breeding

The great progress in super rice breeding both in China and other countries has been made in recent years. However, there were three main problems in super rice breeding: 1) the super rice varieties were still rare; 2) most super rice varieties exhibited narrow adaptability; and 3) current breeding theories emphasized too much on the rice growth model, but they were unpractical in guidance for rice breeding. According to the authors’ experience on the rice breeding, the breeding strategies including three steps (super parent breeding, super hybrid rice breeding and super hybrid rice seed production) were proposed, and the objectives of each step and the key technologies to achieve the goals were elucidated in detail. The super parent of hybrid rice should exhibit excellent performance in all agronomic traits, with the yield or sink capacity reached the level of the hybrid rice control in regional trials. The super hybrid rice combination should meet the following criteria: good rice quality, wide adaptation, lodging resistance, resistance to main insects and diseases, and the yield exceeded above 8% over the control varieties in the national and provincial regional trials. To achieve the goal, the technical strategies, such as selecting optimal combination of the parents, increasing selection pressure, paying more attention to harmony of ideal plant type, excellent physiological traits and all the agronomic traits, should be emphasized. The yield of seed production should reach 3.75 t/ha and 5.25 t/ha for the super hybrid rice combinations derived from early-season and middle-season types of male sterile lines, respectively. The main technologies for raising seed production yield included selecting optimum seed production site, using the male sterile line with large sink capacity and good outcrossing characteristics, and improving the amount of the pollen by intensive cultivation of the male parent. According to the technologies of the three-step breeding on super hybrid rice, two super rice parents, including a male parent 996 and a thermo(photo)-genic male sterile [T(P)GMS] line C815S, were bred. Furthermore, a super early hybrid rice combination, Luliangyou 996, which could be used as a double-season early rice variety in middle and lower reaches of the Yangtze River, China, was bred by using the super rice variety 996 as the male parent, and several hybrid rice combinations with higher yield than control variety in regional trials both of Hunan Province and state were bred with the T(P)GMS line C815S as the female parent.     

Discovery of and Preliminary studies on a Rapid-Leafing Rice Genotype at the Vegetative Growth Stage

The leafing rates of fourteen rice varieties were measured in a sowing-time experiment, and a rapid leafing genotype at the vegetative growth stage was discovered in an indica variety Yanhui 559. The leaf number on the main culm of Yanhui 559 was always 4-5 leaves more than that of Lemont, and the leafing rate of Yanhui 559 was significantly higher than that of Lemont based on similar growth durations from sowing to heading. Furthermore, the difference of the leafing rate was significant at the vegetative growth stage, but not distinctive at the panicle initiation stage. Genetic analysis of the leafing rates in the two backcross populations of Yanhui 559 and Lemont showed that major and quantitative genes controlled the expression of rapid leafing character. Based on results of investigation for some plants with similar growth durations in the backcrossing populations, the rapid leafing genotypes exhibited earlier tillering and more tiller numbers per plant, and its yield components including the number of panicles per plant and number of grains per panicle were superior to those of the slow leafing genotypes. Further research and application feasibility of the rapid leafing genotype in breeding were discussed.     

Effects of Uniconazole on Nitrogen Metabolism and Grain Protein Content of Rice

The effects of uniconazole by soaking seeds and spraying leaves at booting stage with different concentrations （0, 20 and 40 mg/kg） on the nitrogen metabolism of flag leaf and grains after flowering, and rice grain protein content and yield were studied with hybrid rice combination Shanyou 63. Under uniconazole treatment, the soluble protein content in flag leaf was increased in early and middle period of grain filling, but this content was nearly the same as or even lower than that of control at maturity; Glutamine synthetase activity in superior and inferior grains and non-protein nitrogen content in superior grains at early stage of grain development were promoted, and moreover, the transforming speed from non-protein nitrogen to protein nitrogen was accelerated; Non-protein nitrogen content was lower than that of control at maturity, but protein nitrogen content at each stage was higher than those of control; Protein nitrogen content in superior and inferior grains and protein nitrogen absolutely accumulative content in a grain both were enhanced and protein content and yield in rice grain were raised. The application of uniconazole by soaking seeds and spraying leaves raised crude protein content by an average of 7.2% and 8.3%, and protein yield by an average of 13.1% and 13.4%, respectively.     

Molecular Improvement of Grain Weight and Yield in Rice by Using GW6 Gene

Molecular design breeding is one of straightforward approaches to break yield barriers in rice. In this study, GW6 gene for grain length and width from Baodali was transferred into an indica recurrent parent 9311 and a japonica variety Zhonghua 11 （ZH11） using marker-assisted backcross （MAB）. One and three introgression lines were selected for phenotypic analysis from 9311 and ZH11 genetic backgrounds, respectively. SSL-1, an improved 9311 near isogenic line with GW6 performed 11%, 19% and 6.7%higher of grain length, 1000-grain weight and single plant yield, respectively, as compared with 9311. All the three improved ZH11-GW6 lines, R1, R2 and R3, had more than 30% increase in grain weight and about 7%higher in grain yield. Seed plumpness of R1, R2 and R3 was improved synchronously because the three ZH11-GW6 lines contained GIF1 （Grain Incomplete Filling 1）, a dominant grain filling gene. Thus, GW6 has high potential in increasing the yield of inbred lines through MAB, making it an important genetic resource in super hybrid rice breeding. This study provides insights in the utilization of GW6 for large grain and high yield rice breeding via molecular design breeding.     

Parental Selection in Rice Cultivar Improvement

The evaluation of rice (Oryza sativa L.) cultivars assists breeders in identifying useful trait relationships and in selecting parents as donors of specific traits. This study was conducted to compare long-grain rice cultivars using genotype × trait (GT) biplot analysis and determine potential donors of traits related to grain yield and quality. Seventeen cultivars in the 2005 and 2006 Uniform Regional Rice Nursery in Beaumont, Texas, USA were analyzed for 20 traits using GT biplot analysis. The GT biplots showed the diversity among cultivars with regards to yield-related traits. Cultivars recommended as donor parents were: Trenasse, Spring, Presidio, and Cocodrie for high grain yield and head rice rate; Trenasse and Presidio for semi-dwarfness; Banks for tall plant height; Wells for high flag leaf area, panicle weight, and number and mass of filled grains per panicle; Hidalgo for high tiller density; Francis for high number of spikelets per panicle; Spring and Trenasse for short flowering duration; Cheniere for more days to heading and maturity and Spring for less days; and Spring and Hidalgo for high grain weight. Breeders can use these cultivars with specific traits to increase grain yield and quality.     

Changes in Activities of Glutamine Synthetase during Grain Filling and Their Relation to Rice Quality

Four japonica rice varieties differed in cooking and eating qualities were used in a pot experiment to study the relationship between the activities of glutamine synthetase during grain filling and rice quality. The activities of glutamine synthetase gradually increased and then declined as a single peak curve in the course of grain filling. The 15th day after heading was a turning point, before which the enzymatic activities in the inferior rice varieties with high protein content were higher than those in the superior rice varietie with low protein content, and after which it was converse. The activity of glutamine synthetase in grain was correlated with the taste meter value, peak viscosity and breakdown negatively at the early stage of grain filling whereas positively at the middle and late stages. Moreover, it was correlated with the protein content of rice grain and setback positively at the early stage and negatively at the middle and late stages. The correlation degree varied with the course of grain filling. From 15 days to 20 days after heading was a critical stage, in which the direction of correlation between the activity of glutamine synthetase and taste meter value and RVA properties of rice changed.     

Genotypic Differences in Growth and Physiological Responses to Transplanting and Direct Seeding Cultivation in Rice

The field experiments were conducted to investigate the growth and physiological responses of six super hybrid rice combinations to two planting methods,transplanting(TP) and direct seeding(DS) during 2006-2007 and 2007-2008. The 1000-grain weight and number of tillers per plant at the early growth stage,the maximum quantum yield of PSII(Fv/Fm) and transpiration rate(Tr) were higher in DS plants than in TP ones,whereas the grain yield,number of panicles per square meter,seed setting rate,net photosynthetic ...     

Relationships of Ethylene Evolution Rate and 1-Aminocylopropane-1-Carboxylic Acid Concentration in Grains during Filling Period with Appearance Quality of Rice

To elucidate the relationship between ethylene evolution from the grains and the appearance quality of rice,ten different rice genotypes were used to determine the ethylene evolution rate,1-aminocylopropane-1-carboxylic acid(ACC)concentration in grains during grain filling and the appearance quality of rice,and the effects of chemical regulators on concentrations of ethylene and ACC in the grains during grain filling were also investigated to verify the roles of ethylene in the rice quality formation.The ethylene evolution rates and ACC concentrations in grains during the mid and late grain filling stages were very significantly and positively correlated with chalky kernel percentage and chalkiness.The cultivars with a low ACC concentration in grains exhibited a close amyloplast arrangement and small space between starch granules,whereas those with a high ACC concentration in grains showed a loose arrangement and wide space between the granules.Application of 1μmol/L ACC to panicles at mid and late grain filling stages significantly loosened amyloplast arrangement and increased chalky kernel percentage,chalky area and chalkiness,and the results were reversed when 1μmol/L amino-ethoxyvinylglycine,an inhibitor of ACC synthesis enzyme,was applied to panicles.A practice of moderate dry-wet alternate irrigation reduced ethylene evolution and ACC concentration in grains and thereby reduced chalkiness. The results suggested that ethylene and ACC in grains play an important role in the endosperm structure and appearance quality of rice,and the appearance quality would be improved by reducing ethylene evolution and ACC in grains through either variety breeding and selection,or chemical regulations or cultivation techniques.     

The rice miR171b–SCL6-IIs module controls blast resistance,grain yield,and flowering

Micro RNAs(mi RNAs)act as regulators of plant development and multiple stress responses.Here we demonstrate that the rice mi R171 b-SCL6-IIs module regulates the balance between blast resistance,grain yield,and flowering.mi R171 b-overexpressing rice plants(OX171 b)displayed increased rice blast resistance accompanied with enhanced defense responses and late heading,whereas blocking mi R171 b expression in rice(MIM171)led to greater susceptibility to blast disease,associated with compromised defense responses and early heading.Either overexpressing or silencing of mi R171 b significantly affected plant height and number of filled seeds per panicle(seed-setting rate),resulting in decreased grain yield.mi R171 b targets SCL6-IIa,SCL6-IIb,and SCL6-IIc,whose expression was suppressed in OX171 b but increased in MIM171.Mutants of SCL6-IIa,SCL6-IIb,and SCL6-IIc all displayed phenotypes like that of OX171 b,including markedly increased blast disease resistance,slightly decreased grain yield,and delayed flowering.Amounts of mi R171 b increased gradually in leaves during the vegetative stage but decreased gradually in panicles during the reproductive stage,whereas SCL6-IIs displayed the reverse expression pattern.Together,these results suggest that the expression of mi R171 b was time-and space-dependent during the rice growth period and regulated the balance between rice blast disease resistance,grain yield,and flowering via SCL6-IIs,and that appropriate accumulation of mi R171 b is essential for rice development.     

Relationship between grain yield and ^32P, ^14C in tiller and panicle of 56 hybrid rices

In recent years, much progress has been made in China on the relationship between grain yield of F1 hybrid rices and some physiological characters. Experiments were conducted in our plant physiological lab during 1982-1991 to analyze the relationship of the grain yield of F hybrid rice with the distribution and concentration of P and C using isotopes ^32P and ^14C.     

Heterosis Expression of Hybrid Rice in Natural-and Short-Day Length Conditions

Development of hybrid rice is an important strategy to increase yields of irrigated rice in China. Day length is a highly important environmental factor for hybrid rice adaptability,productivity and quality. Heterosis of hybrid rice has been utilized under natural-(long) day length conditions in the middle and lower reaches plain of Yangtze River Valley in China for a long time. However,whether the hybrid rice can show heterosis under short-day length conditions is unknown. In this study,we analyzed plant height,grain number per panicle,and other yield related traits of hybrid rice Liangyoupeijiu(LYP9) and its parents(Pei’ai 64 S and 93-11) under natural- and short-day length conditions and compared the respective heterosis. Results showed that earlier heading,reduced plant height,and reduced grain number per panicle were found in these three varieties under short-day length conditions relative to those under natural-day length conditions. In either natural- or short-day length conditions,heterosis of LYP9 was mainly displayed in grain number per panicle and plant height. This study may help in breeding and selecting suitable hybrid rice for different regions with natural- or short-day length.     

Effect of Grain Position within a Panicle and Variety on As, Cd, Cr, Ni, Pb Concentrations in japonica Rice

Two japonica rice varieties, a compact panicle (CP) Xiushui 63 and a loose panicle (LP) Xiushui 11 were grown in a slightly contaminated paddy field, to determine the effect of grain positions within a panicle and variety on grain As, Cd, Cr, Ni and Pb concentrations. There was a significant difference in grain As, Cd, Cr, Ni and Pb concentrations among grain positions, and the extent of the difference was variety- and metal-specific; Xiushui 63 showed greater difference than Xiushui 11, and As, Cd, Cr and Pb being greater than Ni. The top grains (TG) of a panicle had higher As, Cd and Ni concentrations than the middle ones (MG), and the bottom ones (BG) contained the lowest. With regard to Cr and Pb, the case was opposite. Significantly positive correlations were found between grain weight and As, Cd, Ni concentrations, but there was a negative correlation between grain weight and Cr, and Pb concentrations. There was a remarkable variation in grain filling pattern among grains within a panicle, and between Xiushui 63 and Xiushui 11. In comparison with MG and TG, BG had the lowest grain filling rate and grain weight, leading to obvious difference in grain weight among grains within a panicle, and a greater difference for Xiushui 63 than for Xiushui 11. The regression analysis showed that grain As and Cd concentrations were positively correlated with the maximum grain filling rate (GRm),while negatively correlated with the required time for reaching the maximum grain filling rate (Tpol). Concerning Cr and Pb, the case was just reverse. It is suggested that As and Cd accumulation in grains might be accompanied by the carbohydrate accumulations, while Cr and Pb accumulation displayed a different pattern.     

Preliminary studies on microbiological mechanism of the dynamics of CH4 and N2O emission from rice field

Greenhouse gases, CH4 and N2O emitted from rice field, are the products of microbial metabolism. So the characteristics of CH4 and N2O production and emission in rice field can be understood through microbiological study. In this paper, the relationships between the dynamics of CH4 and N2O emission from rice field in northern China and the related soil bacteria groups were discussed.     

Evaluation of Agronomic Traits in Chromosome Segment Substitution Lines of KDML105 Containing Drought Tolerance QTL under Drought Stress

Drought is a major abiotic constraint to rice production in rainfed lowland and insufficiently irrigated areas.The improvement of drought tolerant varieties is one of the strategies to reduce the negative effects of drought.Quantitative trait loci(QTLs) for primary and secondary traits related to drought tolerance(DT) on chromosomes 1,3,4,8 and 9 that determined from double haploid lines derived from a cross between CT9993 and IR62266 were introgressed and dissected into small pieces in the genetic background of Khao Dawk Mali 105(KDML105) to develop chromosome segment substitution line(CSSL) population.The CSSLs were evaluated at the reproductive stage for their agronomic performance and yield components under drought stress,and results were compared with irrigated condition.The flowering of CSSL lines was 6 to 7 d earlier than KDML105.The mean values of grain yields in the CSSLs were higher than KDML105 under drought and irrigated conditions.At irrigated condition,the grain yields of introgression lines carrying DT-QTLs from chromosomes 4 and 8 were higher than that of KDML105,whereas other traits showed little difference with KDML105.Analysis indicated that grain yield has positive correlation with plant height,tiller and panicle number per plant,and total grain weight per plant under drought stress while negatively correlated with days to flowering.As mentioned above,CSSLs showing good adaptation under drought stress can be used as genetic materials to improve drought tolerance in Thai rainfed lowland rice breeding program,and as materials to dissect genes underlying drought tolerance.     

Solar radiation-use characteristics of indica/japonica hybrid rice (Oryza sativa L.) in the late season in southeast China

New indica and japonica hybrid rice cultivars,such as the Yongyou series,provide farmers with very high yield potential.However,information on their canopy light capture and solar radiation use efficiency in the late season is limited.Field experiments were performed to compare the radiation-use parameters of four rice types:indica rice(IR),inbred japonica rice(IJR),hybrid japonica rice(HJR),and hybrid indica/japonica rice(HIJR),from 2016 to 2018 during the late season in Hangzhou,China.The grain yield,aboveground biomass,intercepted solar radiation(SI),and radiation-use efficiency(RUE)of the HIJR were on average respectively 13.4%–53.4%,14.3%–30.6%,7.6%–21.4%,and 8.2%–14.9%higher than those of the HJR,IJR,and IR.The leaf area index(LAI)of the HIJR was 18.2%–57.0%greater than that of the IJR and HJR at four growth stages,resulting in respectively 17.8%–38.5%and 10.7%–42.8%greater canopy light interception rates(LIR)and amount of intercepted solar radiation during the vegetative stage.The prolonged grain-filling stage also led to respectively 33.9%–52.6%and 30.5%–51.4%increases in amounts of incident and intercepted radiation for the HIJR relative to the IR during grain filling.These results indicate that the SI superiority of the HIJR was caused by canopy closure as rapid as that of the IR during the vegetative stage(greater LAI and canopy LIR during the growing season)and a grain-filling stage as long as that of the HJR.For grain-filling stage,differences in leaf Pn between HIJR,IR,and IJR were not significant,suggesting that the greater RUE of the HIJR(12.7%–52.8%higher)than that of the other rice types resulted from improved canopy architecture after flowering(FL).Principal components analysis(PCA)revealed that the superiority of the HIJR in terms of solar radiation use resulted from the greater canopy light capture capability of IR and the prolonged growth period(especially during grain filling)of japonica rice in the late growing season.