Changes in grain-filling characteristics of single-cross maize hybrids released in China from 1964 to 2014

Grain filling is the physiological process for determining the obtainment of yield in cereal crops.The grain-filling characteristics of 50 maize brand hybrids released from 1964 to 2014 in China were assayed across multiple environments.We found that the grain-filling duration(54.46%)and rate(43.40%)at the effective grain-filling phase greatly contributed to the final performance parameter of 100-kernel weight(HKW).Meanwhile,along with the significant increase in HKW,the accumulated growing degree days(GDDs)for the actual grain-filling period duration(AFPD)among the selected brand hybrids released from the 1960s to the 2010s in China had a decadal increase of 23.41℃ d.However,there was a decadal increase of only 19.76℃ d for GDDs of the days from sowing to physiological maturity(DPM),which was also demonstrated by a continuous decrease in the ratio between the days from sowing to silking(DS)and DPM(i.e.,from 53.24%in the 1960s to 49.78%in the 2010s).In contrast,there were no significant changes in grain-filling rate along with the release years of the selected hybrids.Moreover,the stability of grain-filling characteristics across environments also significantly increased along with the hybrid release years.We also found that the exotic hybrids showed a longer grain-filling duration at the effective grain-filling phase and more stability of the grain-filling characteristics than those of the Chinese local hybrids.According to the results of this study,it is expected that the relatively longer grain-filling duration,shorter DS,higher grain-filling rate,and steady grain-filling characteristics would contribute to the yield improvement of maize hybrids in the future.

Response of grain-filling rate and grain quality of mid-season indica rice to nitrogen application

Nitrogen is one of the important factors for high yield of rice.Apart from high yield,high quality has become the current urgent demand for rice production.Grain-filling stage is crucial for rice yield and quality formation.However,the effect of nitrogen on grain-filling characteristics and the relationship of grain-filling characteristics and rice quality of mid-season indica rice were still unclear.A field experiment was carried out to ascertain the critical grain-filling characteristics that contribute to rice milling quality,appearance quality and cooking and eating quality under nitrogen applications.The results showed that nitrogen applications prolonged the duration of superior and inferior grain filling.The mean grainfilling rate(G_(mean))and the maximum grain-filling rate(G_(max))of the inferior grains were positively correlated with chalky kernel rate,chalkiness,and amylose content.The time reaching the maximum grain-filling rate(T_(max)G)of the inferior grains was positively correlated with brown rice rate,milled rice rate,and head milled rice rate.Chalky kernel rate and chalkiness were negatively correlated with peak paste viscosity and breakdown viscosity.Less amylose content and more crude protein content were detected in nitrogen application of Liangyoupei 9 and Y Liangyou 2 both in 2016 and 2017.According to the correlation analysis,better cooking and eating quality of Y Liangyou 2 which had less amylose content might result from its higher G_(max)and G_(mean)of inferior grain than that of Liangyoupei 9 in the treatments of nitrogen application.These results indicated that the prolonging grain-filling duration and increasing grain weight at the maximum grain-filling rate of inferior grains contributed to the improvement of milling quality,appearance quality,and cooking and eating quality of mid-season indica rice under appropriate nitrogen applications.

Genetic dissection of the grain-filling rate and related traits through linkage analysis and genome-wide association study in bread wheat

Wheat grain yield is generally sink-limited during grain filling.The grain-filling rate(GFR)plays a vital role but is poorly studied due to the difficulty of phenotype surveys.This study explored the grain-filling traits in a recombinant inbred population and wheat collection using two highly saturated genetic maps for linkage analysis and genome-wide association study(GWAS).Seventeen stable additive quantitative trait loci(QTLs)were identified on chromosomes 1B,4B,and 5A.The linkage interval between IWB19555 and IWB56078 showed pleiotropic effects on GFR_(1),GFR_(max),kernel length(KL),kernel width(KW),kernel thickness(KT),and thousand kernel weight(TKW),with the phenotypic variation explained(PVE)ranging from 13.38%(KW)to 33.69%(TKW).198 significant marker-trait associations(MTAs)were distributed across most chromosomes except for 3D and 4D.The major associated sites for GFR included IWB44469(11.27%),IWB8156(12.56%)and IWB24812(14.46%).Linkage analysis suggested that IWB35850,identified through GWAS,was located in approximately the same region as QGFR_(max)2B.3-11,where two high-confidence candidate genes were present.Two important grain weight(GW)-related QTLs colocalized with grain-filling QTLs.The findings contribute to understanding the genetic architecture of the GFR and provide a basic approach to predict candidate genes for grain yield trait QTLs.

Study on the Relationship of the Characteristics of Vascular Bundles in the First Internode and Grain-filling of Heavy Panicle Type Hybrid Rice

The characteristics of vascular bundle in the first internode from top and grain-filling of heavy panicle type hybrid rice(HPTHR) were studied. The results were as follows: The HPTHR had more vascular bundles and a much bigger area of single vascular bundle, all vascular bundle, all phloem and all xylem in the first inter-node than Shanyou63. The vascular bundles had the similar load of spikelet number and sink capacity between the HPTHR and Shanyou63. The HPTHR had not only a larger sink but also normal grain-filling and sink-filling, high seed-setting rate and heavier panicle. Those characteristics of vascular bundle were the biological bases for the larger and heavier panicle of HPTHR. The results also indicated that the breeding model of HPTHR was an effective measure for the super high-yielding rice breeding.

Effect of chemical regulators on the recovery of leaf physiology,dry matter accumulation and translocation,and yield-related characteristics in winter wheat following dry-hot wind

Dry-hot wind stress causes losses in wheat productivity in major growing regions worldwide,especially winter wheat in the Huang-Huai-Hai Plain of China,and both the occurrence and severity of such events are likely to increase with global climate change.To investigate the recovery of physiological functions and yield formation using a new noncommercial chemical regulator(NCR)following dry-hot wind stress,we conducted a three-year field experiment(2018-2021)with sprayed treatments of tap water(control),monopotassium phosphate(CKP),NCR at both the jointing and flowering stages(CFS),and NCR only at the jointing stage(FSJ)or flowering stage(FSF).The leaf physiology,biomass accumulation and translocation,grain-filling process,and yield components in winter wheat were assessed.Among the single spraying treatments,the FSJ treatment was beneficial for the accumulation of dry matter before anthesis,as well as larger increases in the maximum grain-filling rate and mean grain-filling rate.The FSF treatment performed better in maintaining a high relative chlorophyll content as indicated by the SPAD value,and a low rate of excised leaf water loss in flag leaves,promoting dry matter accumulation and the contribution to grain after anthesis,prolonging the duration of grain filling,and causing the period until the maximum grain-filling rate reached earlier.The CFS treatment was better than any other treatments in relieving the effects of dry-hot wind.The exogenous NCR treatments significantly increased grain yields by 12.45-18.20% in 2018-2019,8.89-13.82% in 2019-2020,and 8.10-9.00% in 2020-2021.The conventional measure of the CKP treatment only increased grain yield by 6.69% in 2020-2021.The CFS treatment had the greatest mitigating effect on yield loss under dry-hot wind stress,followed by the FSF and FSJ treatments,and the CKP treatment only had a minimal effect.In summary,the CFS treatment could be used as the main chemical control measure for wheat stress resistance and yield stability in areas with a high incidence of dry-hot wind.This treatment can effectively regulate green retention and the water status of leaves,promote dry matter accumulation and efficient translocation,improve the grain-filling process,and ultimately reduce yield losses.

The spike weight contribution of the photosynthetic area above the upper internode in a winter wheat under different nitrogen and mulching regimes

Besides leaves, non-foliar green organs such as stem and spike are also considered photosynthetic organs. To assess the photosynthetic contributions of these organs, the correlations between these photosynthetic areas and single-spike weight were investigated in a winter wheat(Triticum aestivum L.) under four nitrogen and mulching treatments: N120,N150, N195, and N195 + M. Two-year repeated field experiments were conducted on the Loess Plateau of China. Non-foliar photosynthetic area, grain-filling ratio and duration,grain yield, and in particular, single-spike weight, were measured, recorded and analyzed.Under the N195 + M treatment, plants showed the largest area of photosynthetic organs(flag leaf and non-foliar organs) and the highest grain yield and single spike weight. Singlespike weight was positively correlated with the areas of all examined non-foliar photosynthetic organs, in particular with the area above the flag leaf node(R^2= 0.761*)and the area above the exposed part of the peduncle(EXP)(R^2= 0.800**). In addition, singlespike weight was highly correlated with average grain-filling ratio(R^2= 0.993**), whereas it was less highly correlated with grain-filling duration(R^2= 0.533). The morphological traits of non-foliar photosynthetic organs were also more highly correlated with average grainfilling ratio than with average grain-filling duration. The significant correlation between each of the morphological traits(area, length and width) of EXP and single-spike weight indicates that morphological traits of EXP are important in determining spike weight in the Loess Plateau environment.

Sugar Starvation Enhances Leaf Senescence and Genes Involved in Sugar Signaling Pathways Regulate Early Leaf Senescence in Mutant Rice

To clarify the complex regulatory relationship between changes in sugar content and leaf senescence during the grain-filling stage of rice,genotype-dependent differences in sugar content and the temporal transcriptional patterns of genes involved in sugar signaling pathways were determined in mutant rice exhibiting early leaf senescence and its wild type Zhefu 142.The effects of exogenous glucose or sucrose on the senescence of detached leaves under dark conditions were also investigated.Chlorophyll,soluble sugar,sucrose and fructose contents decreased,whereas electrolytic leakage and malondialdehyde levels increased in mutant leaves at the grain-filling stage.These results suggested that sugar starvation is positively correlated with the early leaf senescence of mutant plants.Detached leaf segments incubated in exogenous sugar solutions under dark conditions exhibited delayed senescence.The high expression of Hxk1 in leaves of mutant plants at the initial grain-filling stage suggested that Hxk1 is involved in the hexose-sensing process at the early stage of leaf senescence.The low expression levels of Hxk2 and Frk1 in the senescing leaves of mutant rice during the grain-filling stage are indicative of weakened hexose phosphorylation.In addition,the high expression levels of Su Sy1,Su Sy2 and Su Sy4 in leaves of mutant plants at the initial grain-filling stage are accompanied by the high transcript levels of SUT1,which favor sucrose translocation and remobilization from the early senescing leaves of mutant rice.The relatively reduced transcript levels of ch FBP,cy FBP,SPS1,SPS2 and SPS6 indicated that during the grain-filling stage,sucrose biosynthesis is weakened in the senescing leaves of mutant rice.

Changes in Starch Accumulation and Activity of Enzymes Associated with Starch Synthesis of Rice at Different N Supplying Dates

The changes in grain-filling, starch accumulation and activity of enzymes associated with starch synthesis in two different hybrid rice varieties were analyzed at different N supplying dates (earlier-date-emphasized, mean-date-emphasized and later-date- emphasized). The results showed that the N application of later-date-emphasized could promote grain-filling rate, increase grain weight and amylopectin content. The peak of activity in three enzymes of ADPglusoce pyriphosphorylase (ADPG), starch synthesis enzyme (SSS) and starch branching enzyme (SBE) in grains of two different rice varieties was not changed obviously, but the mean and maximum activity of these three enzymes changed, and the changes of SSS and ADPG were bigger than that of SBE as N supplying date changed. The N application of earlier-date-emphasized increased SSS activity and the N application of later-date-emphasized increased ADPG and SBE activities. The mean SSS activity during whole grain-filling period, and ADPG and SBE activities at middle and late period of grain-filling were significantly or very significantly correlated with grain-filling rate and accumulating rate of amylose and amylopectin. Both of ADPG and SBE played an equal important role in the changes of amylose and amylopectin content. The N application of later-date-emphasized increased amylose and amylopectin accumulating rate.

The Growth Analysis in the Kernel Filling Process of K-Type Hybrid 901 and Its Parent Wheat

Using Matlab software, the grain filling process of hybrid wheat 901 and its parents was fitted by Richards equation W = A/（1 ＋ Be^-kt in order to study the characteristics of grain filling of the hybrid. The active grain growth period of the hybrid was 6 d longer than that of Shaan 229, and its final grain weight （43.7 g/1 000 grains） was higher than that of Shaan229 （36.3 g/1 000 grains）. N values of 901 and R205 were both less than 1, and their grain growth was faster in the early filling stage, while slower in the middle-late stage. N value of Shaan 229 was 〉1~ and its growth was slower in the early stage and faster in the middle stage. The period of early stage of 901 was shorter and of middle-late stage was longer. The situation of Shaan 229 was totally reversed. For parents, the father plant R205 was similar to hybrid wheat 901, whereas its mother plant K3314A similar to Shaan 229. It has been found that Richards equation was more suitable for fitting the grain filling process of wheat than Logistic equation.

Causes of Poor Grain Plumpness of Two-Line Hybrids and Their Relationships to the Contents of Hormones in the Rice Grain

This study was designed to elucidate the grain filling characteristics and the causes of poor grain plumpness of some two-line hybrid rice combinations and their hormonal mechanism. Six varieties, including three two-line hybrid rice combinations, that show differences in seed-setting and grain filling, were used. And the contents of starch, sucrose, zeatin （Z） ＋ zeatin riboside （ZR）, indole-3-acetic acid （IAA）, and abscisic acid （ABA）, the ethylene evolution rate, activities of sucrose synthase （SuSase） and starch synthase （StSase） in grains, the seed-setting and grain filling rate were investigated. The correlations amongst these were analyzed. The results showed that the poor grain filling of two-line hybrids was mainly attributed to the higher unfilled grain rate and the lower filling degree of inferior grains. During the early and mid grain filling periods, the sucrose content in inferior grains was greater than that in superior grains for the combinations with poor grain filling, indicating that the substrate concentration was not the principal factor for their slow grain filling and poor grain plumpness of the inferior grains of two-line hybrids. Z ＋ ZR, IAA, and ABA in superior grains were obviously greater than those in inferior grains at early grain filling stage. The maximum and mean contents of Z ＋ ZR, IAA, and ABA were positively very significantly correlated with the maximum and mean grain-filling rate, filling degree, and grain weight. The evolution rate of ethylene was greater in inferior grains than in superior grains and greater for the combinations with poor grain plumpness than those with good grain plumpness at the early or mid filling stages. The evolution rate of ethylene was negatively and significantly correlated with the grain filling rate, the grain filling degree, and the grain weight. Spraying ethephon （ethylene-releasing agent） at the early grain filling stage increased the evolution rate of ethylene, reduced the ABA content and activities of SuSase and StSase, and decreased the grain filling degree and the grain weight. The results were reversed when cobatous nitrate （an inhibitor of ethylene synthesis） was applied. The results suggested that the hormones and their balance play a role in the regulation of grain filling and enzymatic activities, and the poor grain filling is attributed to the low contents of Z ＋ ZR, IAA, and ABA, and the high evolution rate of ethylene in the inferior grains of some two-line hybrid rice combinations. The results suggested that hormones play important roles in the grain filling of some two-line hybrid rice combinations, and their filling degree can be improved by regulating the hormonal contents.

Identification of Differentially Senescing Mutants of Wheat and Impacts on Yield,Biomass and Nitrogen Partitioning

Increasing photosynthetic capacity by extending canopy longevity during grain filling using slow senescing stay-green genotypes is a possible means to improve yield in wheat. Ethyl methanesulfonate （EMS） mutated wheat lines （Triticum aestivum L. cv. Paragon） were screened for fast and slow canopy senescence to investigate the impact on yield and nitrogen partitioning. Stay-green and fast-senescing lines with similar anthesis dates were characterised in detail. Delayed senescence was only apparent at higher nitrogen supply with low nitrogen supply enhancing the rate of senescence in all lines. In the stay-green line 3 （SG3）, on a whole plant basis, tiller and seed number increased whilst thousand grain weight （TGW） decreased; although a greater N uptake was observed in the main tiller, yield was not affected. In fast-senescing line 2 （FS2）, yield decreased, principally as a result of decreased TGW. Analysis of N-partitioning in the main stem indicated that although the slow-senescing line had lower biomass and consequently less nitrogen in all plant parts, the proportion of biomass and nitrogen in the flag leaf was greater at anthesis compared to the other lines; this contributed to the grain N and yield of the slow-senescing line at maturity in both the main tiller and in the whole plant. A field trial confirmed senescence patterns of the two lines, and the negative impact on yield for FS2 and a positive impact for SG3 at low N only. The lack of increased yield in the slow-senescing line was likely due to decreased biomass and additionally a possible sink limitation.