Performance in Grain Yield and Physiological Traits of Rice in the Yangtze River Basin of China During the Last 60 yr

Knowledge on the performance in grain yield and physiological traits is essential to understand the main yield-limiting factor and make strategies for breeding and crop management in rice （Oryza sativa L.）. This study investigated the changes in grain yield and associated physiological traits of rice in the Yangtze River Basin of China during the last 60 yr. Thirteen mid-season indica and 12 japonica rice cultivars that were popularly used were grown in the field in 2008 and 2009. The grain yield and yield components, biomass, leaf area, leaf photosynthesis, root oxidation activity, and harvest index were examined. The results showed that grain yield and grain yield per day have progressively increased during the years and such increases are mainly attributed to the expanded sink size as a result of more spikelets per panicle, especially for the case of super rice. Both biomass and harvest index were increased with the improvement of cultivars. Increase in biomass for modern rice cultivars was associated with an enhancement of leaf area and photosynthesis, root dry weight, and root oxidation activity, although the indica super rice cultivars showed a lower leaf photosynthetic rate and root oxidation activity than the semi-dwarf cultivars during the grain filling period. Both indica and japonica super rice cultivars exhibited a low percentage of filled grains, which may limit their great yield potential. All the data suggested that grain yield have been substantially improved during the 60 yr of rice breeding in the Yangtze River Basin. Expanded sink size, increased dry matter production and harvest index, and enhanced leaf area and photosynthesis, root dry weight, and root oxidation activity contribute to the improvement in grain yield. Increase in filling efficiency could realize the great yield potential in super rice.

Characterization and mapping of QTLs on chromosome 2D for grain size and yield traits using a mutant line induced by EMS in wheat

Production of mutants with altered phenotypes is a powerful approach for determining the biological functions of genes in an organism. In this study, a high-grain-weight mutant line M8008 was identified from a library of mutants of the common wheat cultivar YN15 treated with ethylmethane sulfonate(EMS). F2 and F2:3generations produced from crosses of M8008 × YN15(MY) and M8008 × SJZ54(MS) were used for genetic analysis. There were significant differences between M8008 and YN15 in plant height(PH), spike length(SL),fertile spikelet number per spike(FSS), grain width(GW), grain length(GL), GL/GW ratio(GLW), and thousand-grain weight(TGW). Most simple correlation coefficients were significant for the investigated traits, suggesting that the correlative mutations occurred in M8008. Approximately 21% of simple sequence repeat(SSR) markers showed polymorphisms between M8008 and YN15, indicating that EMS can induce a large number of mutated loci. Twelve quantitative trait loci(QTLs) forming QTL clusters(one in MY and two in MS) were detected. The QTL clusters coinciding with(MY population) or near(MS population) the marker wmc41 were associated mainly with grain-size traits, among which the M8008 locus led to decreases in GW, factor form density(FFD), and TGW and to increases in GLW. The cluster in the wmc25–barc168 interval in the MS population was associated with yield traits, for which the M8008 locus led to decreased PH, spike number per plant(SN), and SL.

Relationship Between Heterosis and Parental Genetic Distance Based on Molecular Markers for Functional Genes Related to Yield Traits in Rice

The genetic distances among 18 cytoplasmic male sterile lines and 11 restorer lines were analyzed with molecular markers derived from yield-related functional genes. The correlation between parental genetic distance and heterosis was investigated by analyzing the performance of 47 combinations. The results showed that the genetic distance was significantly correlated with yield heterosis （r=0.29^＊）, but not significantly correlated with heterosis for other traits, such as number of effective panicles per plant, seed setting rate, 1000-grain weight, number of grains per panicle and theoretical yield. However, the correlation coefficient was so small that the parental genetic distance could not to be used to predict heterosis.

Dissection of heterosis for yield and related traits using populations derived from introgression lines in rice

Despite the great success achieved by the exploitation of heterosis in rice,the genetic basis of heterosis is still not well understood.We adopted an advanced-backcross breeding strategy to dissect the genetic basis of heterosis for yield and eight related traits.Four testcross(TC) populations with 228 testcross F1 combinations were developed by crossing57 introgression lines with four types of widely used male sterile lines using a North Carolina II mating design.Analysis of variance indicated that the effects of testcross F1 combinations and their parents were significant or highly significant for most of the traits in both years,and all interaction effects with year were significant for most of the traits.Positive midparent heterosis(HMP) was observed for most traits in the four TC populations in the two years.The relative HMPlevels for most traits varied from highly negative to highly positive.Sixty-two dominant-effect QTL were identified for HMPof the nine traits in the four TC populations in the two years.Of these,22 QTL were also identified for the performance of testcross F1.Most dominant-effect QTL could individually explain more than 10% of the phenotypic variation.Four QTL clusters were observed including the region surrounding the RM9–RM297 region on chromosome 1,the RM110–RM279–RM8–RM5699–RM452 region on chromosome 2,the RM5463 locus on chromosome 6 and the RM1146–RM147 region on chromosome 10.The identified QTL for heterosis provide valuable information for dissecting the genetic basis of heterosis.

Identification of QTLs for Yield-Related Traits in the Recombinant Inbred Line Population Derived from the Cross Between a Synthetic Hexaploid Wheat-Derived Variety Chuanmai 42 and a Chinese Elite Variety Chuannong 16

Synthetic hexaploid wheat （SHW） represents a valuable source of new resistances to a range of biotic and abiotic stresses. A recombinant inbred line （RIL） population with 127 recombinant inbred lines derived from a SHW-derived variety Chuanmai 42 crossing with a Chinese spring wheat variety Chuannong 16 was used to map QTLs for agronomic traits including grain yield, grains per square meter, thousand-kernel weight, spikes per square meter, grain number per spike, grains weight per spike, and biomass yield. The population was genotyped using 184 simple-sequence repeat （SSR） markers and 34 sequence-related amplified polymorphism （SRAP） markers. Of 76 QTLs （LOD〉2.5） identified, 42 were found to have a positive effect from Chuanmai 42. The QTL QGy.saas-4D.2 associated with grain yield on chromosome 4D was detected in four of the six environments and the combined analysis, and the mean yield, across six environments, of individuals carrying the Chuanmai 42 allele at this locus was 8.9% higher than that of those lines carrying the Chuannong 16 allele. Seven clusters of the yield-coincident QTLs were detected on 1A, 4A, 3B, 5B, 4D, and 7D.

Response of phenology- and yield-related traits of maize to elevated temperature in a temperate region

Extreme high temperatures detrimental to maize production are projected to occur more frequently with future climate change.Phenology and yield-related traits were investigated under several levels of elevated temperature in two early-maturing hybrid cultivars:Junda 6(grown in northeastern China)and Chalok 1(grown in South Korea).They were cultivated in plastic houses in Suwon,Korea(37.27°N,126.99°E)held at target temperatures of ambient(AT),AT+1.5°C,AT+3°C,and AT+5°C at one sowing date in 2013 and three different sowing dates in 2014.Vegetative and reproductive growth durations showed variation depending on sowing date,experimental year,and cultivar.Growth duration tended to decrease,but not necessarily,with temperature elevation,but somewhat increased again above a certain temperature.High temperature-dependent variation was greater during grain filling than in the vegetative period before anthesis.Elevated temperature showed no significant effects on duration or peak dates of silking and anthesis,and thus on anthesis–silking interval.Grain yield tended to decrease with temperature elevation above ambient,showing a sharper linear decrease with mean growing season temperature increase in Junda 6 than in Chalok 1.The decrease in kernel number accounted for a much greater contribution to the yield reductions due to temperature elevation than did the decrease in individual kernel weight in both cultivars.Individual harvestable kernel weight was not significantly affected by temperature elevation treatments.Kernel number showed a linear decrease with mean growth temperature from early ear formation to early grain-filling stage,with Junda 6 showing a much severer decrease than Chalok 1.Kernel number reduction due to temperature elevation was attributable more to the decrease in differentiated ovule number than to the decrease in kernel set in Chalok 1,but largely to the decrease of kernel set in Junda 6.

Major Gene Identification and Quantitative Trait Locus Mapping for YieldRelated Traits in Upland Cotton(Gossypium hirsutum L.)

Segregation analysis of the mixed genetic model of major gene plus polygene was used to identify the major genes for cotton yield-related traits using six generations P1, P2, F1, B1, B2, and F2 generated from the cross of Baimian 1 x TM-1. In addition to boll size and seed index, the major genes for the other five traits were detected： one each for seed yield, lint percentage, boll number, lint index; and two for lint yield. Quantitative trait locus/loci （QTL） mapping was performed in the F2 and F2：3 populations of above cross through molecular marker technology, and a total of 50 QTL （26 suggestive and 24 significant） for yield-related traits were detected. Four common QTL were discovered： qLP-3b（F2）/qLP-3（F2：3） and qLP-19b （F2）/qLP-19（F2：3） for lint percentage, qBN-17（F2）/qBN-17（F2：3） for boll number, and qBS-26b（F2）/qBS-26（F2：3） for boll size. Especially, qLP- 3b（Fz）/qLP-3（F2：3）, not only had LOD scores 〉3 but also exceeded the permutation threshold （5.13 and 5.29, respectively）, correspondingly explaining 23.47 and 29.55% of phenotypic variation. This QTL should be considered preferentially in marker assisted selection （MAS）. Segregation analysis and QTL mapping could mutually complement and verify, which provides a theoretical basis for genetic improvement of cotton yield-related traits by using major genes （QTL）.

Detection of Favorable Alleles for Quality- and Yield-related Traits in Wheat Using a Backcross Population

Development of wheat varieties with high yield and good quality has been a major objective in wheat breeding.A BC 1 F 2-3 population was used to detect QTLs for wheat quality related traits: SDS-sedimentation value (Ssd),grain protein content (GPC),grain hardness (GH) and 11 mixograph parameters,as well as five agronomic traits: spike length (SL),spikelet number per spike (SPN),grain number per spike (GN),thousand-grain weight (TGW),and plant height (PH).A total of 44 putative QTLs were detected in the present study,31 for quality parameters and 13 for important agronomic traits,including three important major QTLs.One major QTL for Ssd QSsd.saas-1B.1,linked to barc137,explained on average 21.1% of the phenotypic variation in three environments.The allele increasing Ssd at this locus also significantly increased GN.The second locus on chromosome 1B with the linked marker Barc 61 was a major locus for mixograph parameters.It explained 21.3%-32.5%,24.3%-30.6%,30.6%-37% and 20.1%-22.7% of phenotypic variation for mixing tolerance (MT),weakening slope (WS),midline peak time (MPTi) and midline time x =8 value (MTxW),respectively.The third major QTL,explaining above 40% of plant height variation,close to Rht-B 1 on the short arm of chromosome 4BS,co-located with QTL for quality and yield-related traits.

Quantitative trait loci associated to agronomic traits and yield components in a Sorghum bicolor L.Moench RIL population cultivated under pre-flowering drought and well-watered conditions

The present study aims to identify QTL influencing agronomic traits and yield components under well-watered and pre-flowering drought stress conditions. One hundred F5 recombinant inbred lines (RIL) and the parental lines of a cross between a drought-tolerant and a susceptible line in a field experiment were carried out at Nong Lam University of Ho Chi Minh City, Vietnam. Drought stress was induced by withholding irrigation water from the plants at four weeks after sowing to flowering. Leaf area of the third leaf, stem diameter, plant height, days to heading, anthesis and maturity, panicle length, number of seeds per plant, hundred kernel weight and grain yield were measured. Plants were genotyped with 117 Diversity Arrays Technology (DArT) and eight expressed sequence tag (EST)-derived simple sequence repeat (SSR) markers. Composite interval mapping was carried out on the traits and significant QTL were claimed at a logarithm of the odds (LOD) score >2.5. A total of 50 QTL were detected on nine chromosomes or 13 linkage groups, respectively. Six promising QTL regions with seven QTL for yield and agronomic traits especially related to pre-flowering drought tolerance were identified on chromosomes SBI-01, SBI-03, SBI-04, SBI-05 and SBI-07.

Root morphological traits and their relations with yield components of dry-raised mid-season indica rice

Research on root morphological traits of dry-raised seedlings (D-RS) at different growth stages of rice have so far attracted less attention. In this study, using mid-season indica hy-

Lodging Resistance Related to Root Traits for Mechanized Wet-Seeding of Two Super Rice Cultivars

Mechanical hill wet-seeded rice machine is beneficial for establishing and growing uniform rows of seedlings.However,there is limited knowledge regarding the effects of the establishment of furrows on growth,lodging and yield,and their relationships with root traits.In this study,field experiments were conducted during 2012 and 2013 using two super rice varieties(hybrid rice Peizataifeng and inbred rice Yuxiangyouzhan)under three furrow establishment treatments(T1,both water and seed furrows were established by the machine;T2,only seed furrows were established by the machine;and T3,neither water nor seed furrows were established by the machine).Lodging index,lodging-related traits,grain yield,above-ground dry weight and root traits were measured.The results showed that the lodging index was significantly affected by the treatments with furrows(T1 and T2).The strongest lodging resistance was detected in the mechanical hill wet-seeded rice with furrow treatment(T1)in both 2012 and 2013.Lodging resistance was strongly related to the breaking resistance.No significant difference was found in grain yield or dry weight of the mechanical hill wet-seeded rice.Therefore,the mechanical hill wet-seeded rice with furrow treatment increased rice lodging resistance,which was related to root traits.

QTL for grain yield and yield parameters in winter wheat(Triticum aestivum L.)

Field trials with a set of 108 doubled haploid lines(DHs) derived from a cross between the Chinese winter wheat cvs.CA9613 and H1488 were run at Beijing(China).Phenotypic data were recorded for major agronomic yield traits,i.e.grain weight per ear,grain number per ear and thousand grain weight(Tgw) in two field trials at Beijing.Based on the phenotypic data and a genetic map comprising 168 SSR markers,an analysis of quantitative trait loci(QTL) was carried out for yield and yield parameters using the composite interval mapping(CIM) approach.A total of 14 QTL were detected for these traits across two environments.Four of these QTL located on chromosomes 1A and 2B,respectively,exhibited pleiotropic effects.Loci showing pleiotropic effects will be very useful for understanding the homeologous relationships of QTL and designing an appropriate marker-assisted selection programme by multi-trait selection in order to accumulate("pyramide") favorable alleles at different loci.

Optimal yield-related attributes of irrigated rice for high yield potential based on path analysis and stability analysis

Improvement of yield in rice(Oryza sativa L.) is vital for ensuring food security in China. Both rice breeders and growers need an improved understanding of the relationship between yield and yield-related traits. New indica cultivars(53 in 2007 and 48 in 2008) were grown in Taoyuan,Yunnan province, to identify important components contributing to yield. Additionally, two standard indica rice cultivars with similar yield potentials, II You 107(a large-panicle type) and Xieyou 107(a heavy-panicle type), were planted in Taoyuan, Yunnan province and Nanjing,Jiangsu province, from 2006 to 2008 to evaluate the stability of yield and yield-related attributes.Growth duration(GD), leaf area index(LAI), panicles per m2(PN), and spikelets per m2(SM) were significantly and positively correlated with grain yield(GY) over all years. Sequential path analysis identified PN and panicle weight(PW) as important first-order traits that influenced grain yield. All direct effects were significant, as indicated by bootstrap analysis. Yield potential varied greatly across locations but not across years. Plant height(PH), days from heading to maturity(HM), and grain weight(GW) were stable traits that showed little variation across sites or years, whereas GD(mainly the pre-heading period, PHP) and PN varied significantly across locations. To achieve a yield of 15 t ha-1, a cultivar should have a PH of 110–125 cm, a long GD with HM of approximately 40 days, a PN of 300–400 m-2, and a GW of 29–31 mg.

Assessment of molecular markers and marker-assisted selection for drought tolerance in barley(Hordeum vulgare L.)

This review updates the present status of the field of molecular markers and marker-assisted selection(MAS),using the example of drought tolerance in barley.The accuracy of selected quantitative trait loci(QTLs),candidate genes and suggested markers was assessed in the barley genome cv.Morex.Six common strategies are described for molecular marker development,candidate gene identification and verification,and their possible applications in MAS to improve the grain yield and yield components in barley under drought stress.These strategies are based on the following five principles:(1)Molecular markers are designated as genomic‘tags’,and their‘prediction’is strongly dependent on their distance from a candidate gene on genetic or physical maps;(2)plants react differently under favourable and stressful conditions or depending on their stage of development;(3)each candidate gene must be verified by confirming its expression in the relevant conditions,e.g.,drought;(4)the molecular marker identified must be validated for MAS for tolerance to drought stress and improved grain yield;and(5)the small number of molecular markers realized for MAS in breeding,from among the many studies targeting candidate genes,can be explained by the complex nature of drought stress,and multiple stress-responsive genes in each barley genotype that are expressed differentially depending on many other factors.

Comparison of Grain Quality Characteristics Between F_1 Hybrids and Their Parents in Indica Hybrid Rice

Sixteen widespread elite indica parents including seven CMS lines and nine restorer lines, and their 63 F_1 hybrids were chosen to compare the eleven traits of grain quality. Overall results showed that the frequency of negative over-dominance, dominance and partial dominance was much higher than that of positive ones and heterosis, indicating that the values of F_1 hybrids were generally lower than the means of their parents in quality characteristics. There existed apparent disparity in grain quality performance among F_1 hybrids varied with the traits. The characteristics of chalky area percentage, chalky grain percentage, gel consistency and head rice recovery had a great variation in all kinds of tested heterosis indices including the mid-parent heterosis index, over high-value parent heterosis index and over low-value parent heterosis index, which suggested that special attention should be paid to the selection of F_1 hybrids in these traits in breeding. The mean values of F_1 hybrids were significantly lower than those of their higher parents in all the traits but close to or significantly lower than those of their mid-parent values except for kernel length and amylose content; and significantly higher than those of their lower parents except for brown rice recovery, milled rice recovery and gel consistency.

Increased grain yield with improved photosynthetic characters in modern maize parental lines

The grain yield of maize has increased continuously in past decades, largely through hybrid innovation, cultivation tech-nology, and in particular, recent genetic improvements in photosynthesis. Elite inbred lines are crucial for innovating new germplasm. Here, we analyzed variations in grain yield and a series of eco-physiological photosynthetic traits after anthesis in sixteen parental lines of maize (Zea mays L.) released during three different eras (1960s, 1980s, 2000s). We found that grain yield and biomass signiifcantly increased in the 2000s than those in the 1980s and 1960s. Leaf area, chlorophyl , and soluble protein content slowly decreased, and maintained a higher net photosynthesis rate (Pn) and improved stomatal conductance (Gs) after anthesis in the 2000s. In addition, the parental lines in the 2000s obtained higher actual photo-chemistry efifciency (ФPSI ) and the maximum PSII photochemistry efifciency (Fv/Fm), which largely improved light partition-ing and chlorophyl lfuorescence characteristic, including higher photochemical and photosystem II (PSII) reaction center activity, lower thermal energy dissipation in antenna proteins. Meanwhile, more lamel ae per granum within chloroplasts were observed in the parental lines of the 2000s, with a clear and complete chloroplast membrane, which wil greatly help to improve photosynthetic capacity and energy efifciency of ear leaf in maize parental lines. It is concluded that grain yield increase in modern maize parental lines is mainly attributed to the improved chloroplast structure and more light energy catched for the photochemical reaction, thus having a better stay-green characteristic and stronger photosynthetic capac-ity after anthesis. Our direct physiological evaluation of these inbred lines provides important information for the further development of promising maize cultivars.

Co-planted barnyardgrass reduces rice yield by inhibiting plant above-and belowground-growth during post-heading stages

Barnyardgrass (Echinochloa spp.) is the most common noxious weed in rice paddies as it inhibits rice growth and reduces grain yield.To date,little information is available on above-and belowgroundgrowth changes in rice due to neighboring barnyardgrass.This study aimed to investigate the changes in root traits and shoot growth of rice when it is grown with different kinds of barnyardgrass.Japonica rice plants (var.Nanjing 9108) were co-cultured with two varieties of Echinochloa crusgalli (L.) Beauv.(EP,var.mitis (pursh) Petern;EH,var.zelayensis (H.B.K.) Hitchc),and E.colonum (L.) Link (EL) in the field in 2017 and 2018.Four treatments included control (i.e.,weed free rice plants) and co-cultures with each of three barnyardgrasses (EP,EH,and EL).The results revealed that EP,EH,and EL treatments significantly reduced rice grain yields by 30.6%–36.2%,42.5%–46.5%,and 10.6%–14.3%,respectively.Shoot growth including shoot dry weight,leaf photosynthetic rate,zeatin (Z) and zeatin riboside (ZR) in grains,and activities of key enzymes involved in sucrose-to-starch conversion in grains and root traits,such as length density,root dry weight,total absorbing surface area,active absorption surface area,oxidation activity,and Z+ZR contents in roots were dramatically reduced during post-heading stages of rice when grown with the three kinds of barnyardgrass.Moreover,above-mentioned rice shoot growth indices were strongly and positively correlated with root traits.These results suggested the decrease in rice shoot growth and root traits during post-heading stages contributes to the reduction in the rice yield when it grows with barnyardgrass neighbors.

Temporal and spatial variations of carbon isotope signature reveal substantial contribution of bracts and internode assimilates to grain filling of japonica rice

Carbon isotope composition(δ^(13)C)of a plant organ is an inherent signature reflecting its physiological property,and thus is used as an integrative index in crop breeding.It is also a non-intrusive method for quantifying the relative contribution of different source organs to grain filling in cereals.Using the samples collected from two-year field and pot experiments with two nitrogen(N)fertilization treatments,we investigated the temporal and spatial variations of δ^(13)C in source organs of leaf,sheath,internode,and bracts,and in sink organ grain.Constitutive nature of δ^(13)C was uncovered,with an order of leaf(−27.84‰)<grain(−27.82‰)<sheath(−27.24‰)<bracts(−26.81‰)<internode(−25.67‰).For different positions of individual organs within the plant,δ^(13)C of the leaf and sheath presented a diminishing trend from the top(flag leaf and its sheath)to the bottom(the last leaf in reverse order and its sheath).No obvious pattern was found for the internode.For temporal variations, δ^(13)C of the leaf and sheath had a peak(the most negative)at 10 days after anthesis(DAA),whereas that of the bracts showed a marked increase at the time point of anthesis,implying a transformation from sink to source organ.By comparing the δ^(13)C in its natural abundance in the water-soluble fractions of the sheath,internode,and bracts with the δ^(13)C in mature grains,the relative contribution of these organs to grain filling was assessed.With reference to the leaf,the internode accounted for as high as 32.64%and 42.56%at 10 DAA and 20 DAA,respectively.Meanwhile,bracts presented a larger contribution than the internode,with superior bracts being higher than inferior bracts.In addition,N topdressing reduced the contribution of the internode and bracts.Our findings clearly proved the actual significance of non-foliar organs of the internode and bracts for rice yield formation,thus extending our basic knowledge of source and sink relations.

Evaluating Rice Biodiversity and Yields of Upland Rice Landraces Grown in Shifting Cultivation in Bandarban,Bangladesh

Shifting cultivation,popularly known as jhum,is a dominant form of agriculture in the Chattogram Hill Tracts(CHT)of Bangladesh with upland rice being the major component of the system.The region is known for its rice biodiversity,which is under threat.This study was an attempt to explore the extent of rice biodiversity and variation in rice yields observing 81 randomly selected shifting cultivation plots from 26 dispersedly located mountainous villages in four sub-districts of Bandarban,one of three districts of the CHT.A total of 28 landraces of upland rice was grown in shifting cultivation.Highest number of landraces(16)was found in sub-district Thanchi.Three landraces most frequently observed were Gunda,Maemonsing and Sadabinni.Rice grain yield varied between 1.421 t ha-1 and 3.442 t ha-1 across landraces with the highest being recorded for Patobi.Landrace Dilon the lowest yield.Relative performance of landraces Kobrokbinni,Maemonsing,Monthon,Patobi and PD were superior to standard BRRI dhan83 and Gunda in relation to grain yield.Some of these landraces having wider adaptability may be released as varieties.