Analysis of Wheat Spike Morphological Traits by 2D Imaging

BackgroundGrain number (GN) is one of the key yield contributing factors in modern wheat (Triticum aestivum) varieties. Fruiting efficiency (FE) is a key trait for increasing GN by making more spike assimilates available to reproductive structures. Thousand grain weight (TGW) is also an important component of grain yield. To understand the genetic architecture of FE and TGW, we performed a genome-wide association study (GWAS) in a panel of 236 US soft facultative wheats that were phenotyped in three experiments at two locations in Florida and genotyped with 20,706 single nucleotide polymorphisms (SNPs) generated from genotyping-by-sequencing (GBS).ResultsFE showed significant positive associations with GN, grain yield (GY), and harvest index (HI). Likewise, TGW mostly had a positive correlation with GY and HI, but a negative correlation with GN. Eighteen marker-trait associations (MTAs) for FE and TGW were identified on 11 chromosomes, with nine MTAs within genes. Several MTAs associated with other traits were found within genes with different biological and metabolic functions including nuclear pore complex protein, F-box protein, oligopeptide transporter, and glycoside vacuolar protein. Two KASP markers showed significant mean differences for FE and TGW traits in a validation population.ConclusionsKASP marker development and validation demonstrated the utility of these markers for improving FE and TGW in breeding programs. The results suggest that optimizing intra-spike partitioning and utilizing marker-assisted selection (MAS) can enhance GY and HI.

Grain number and weight within a spikelet are major yield components which determine the grain yield in wheat. The objective of this study was to explore genetic gains in grain performance within wheat spikelets at the individual grain level and its effect on grain yield and evaluate genetic progress in stem internode length and other yield-related traits. We conducted field experiments across three growing seasons in the western Yellow and Huai Valley of China; incorporating 17 bread wheat cultivars released from 1948 to 2012. Yields were significantly correlated with year of release. Yield gains equated to 3.95 g m−2 yr−1 in response to increases in total grain weight per m2 (GW) in proximal (G1 and G2) and distal (G3 and G4) grains, despite a decline in the proportional contribution of proximal grains to yield and increase in the proportional contribution of distal grains to yield with year of release. Grain number per m2 (GN), thousand-grain weight (TGW), and harvest index (HI) increased with year of release, but plant height decreased. Both grain number per spikelet of proximal and distal grain contributed to the increase in total GN. However, the contribution ratio of GN in proximal grains to total GN declined, and the proportion in distal grains increased. Average single grain weight (SGW) increased linearly at G1, G2, G3, and G4 with year of release and contributed to the increase in TGW. The G3 and G4 grain positions had much lower individual grain weights but increased at a faster rate than G1 and G2. At G1, G2, and G3 grain positions, from bottom to top spikelets, the newly released cultivars had the heaviest grains and the old cultivars had the lightest grains. New cultivars had more spikelets than old cultivars and the number of grains (proximal and distal grains) in the apical spike increased with year of release. The length of five internodes decreased significantly with year of release, more so in the upper than lower internodes, all of which contributed to the decline in plant height. In summary, increasing the number and weight of distal grains could increase grain yield, TGW, and GN.

Growth, yield, and nitrogen use in the wheat/maize intercropping system in an arid region of northwestern China

Identification of a pleiotropic QTL cluster for Fusarium head blight resistance, spikelet compactness, grain number per spike and thousand-grain weight in common wheat

Research progress on spike development of wheat.

Manipulation of genes involved in starch synthesis could significantly affect wheat grain weight and yield. The starch-branching enzyme (SBE) catalyzes the formation of branch points by cleaving the α-1,4 linkage in polyglucans and reattaching the chain via an α-1,6 linkage. Three types of SBE isoforms (SBEI, SBEII, and SBEIII) exist in higher plants, with the number of SBE isoforms being species-specific. In this study, the coding sequence of the wheat TaSBEIII gene was amplified. After the multiple sequence alignment of TaSBEIII genome from 20 accessions in a wheat diversity panel, one SNP was observed in TaSBEIII-A, which formed the allelic marker allele-T. Based on this SNP at 294 bp (C/T), a KASP molecular marker was developed to distinguish allelic variation among the wheat genotypes for thousand grain weight (TGW). The results were validated using 262 accessions of mini core collection (MCC) from China, 153 from Pakistan, 53 from CIMMYT, and 17 diploid and 18 tetraploid genotypes. Association analysis between TaSBEIII-A allelic variation and agronomic traits found that TaSBEIII-A was associated with TGW in mini core collection of China (MCC). The accessions possessing Allele-T had higher TGW than those possessing Allele-C; thus, Allele-T was a favorable allelic variation. By analyzing the frequency of the favorable allelic variation Allele-T in MCC, it increased from pre-1950 (25%) to the 1960s (45%) and increased continuously from 1960 to 1990 (80%). The results suggested that the KASP markers can be utilized in grain weight improvement, which ultimately improves wheat yield by marker-assisted selection in wheat breeding. The favorable allelic variation allele-T should be valuable in enhancing grain yield by improving the source and sink simultaneously. Furthermore, the newly developed KASP marker validated in different genetic backgrounds could be integrated into a breeding kit for screening high TGW wheat.

Multiple studies have identified genes affecting grain morphology, yet their capacity to deliver yield gains under field conditions remains unclear. We performed a multiyear, multilocation factorial evaluation of GRAIN WIDTH2 ( TaGW2 ) mutants in hexaploid wheat using BC 4 near-isogenic lines, sowing-density treatments and semi-dwarfing RHT1 backgrounds. Loss-of-function mutations in TaGW2 increased grain size and thousand grain weight (TGW) additively; with the aaBBDD single mutant showing the most stable singlelocus effect, while the aabbdd triple mutant achieved ~20% higher TGW across twelve field trials. However, overall grain yield remained unchanged or slightly reduced, reflecting a compensatory trade-off with grain number. Spike phenotyping of both main and secondary tillers showed comparable increases in TGW and spike yield despite fewer grains per spike, indicating that limited yield gain primarily reflects reduced spike number per unit area rather than decreased spike-level productivity. Effects were stable across sowing densities, whereas interactions with semi-dwarfing alleles were allele-specific: RHT-B1b partially suppressed TGW gains and accentuated yield penalties, whereas RHT-D1b maintained the large-grain phenotype and productivity. Across experiments, the TaGW2-A1D1 double mutant increased TGW (~14%) while maintaining yield stability, identifying it as a promising genotype for breeding. We conclude that TaGW2 is a reliable modifier of grain size but not yield in isolation.

BackgroundYield-related traits including thousand grain weight (TGW), grain number per spike (GNS), grain width (GW), grain length (GL), plant height (PH), spike length (SL), and spikelet number per spike (SNS) are greatly associated with grain yield of wheat (Triticum aestivum L.). To detect quantitative trait loci (QTL) associated with them, 193 recombinant inbred lines derived from two elite winter wheat varieties Chuanmai42 and Chuanmai39 were employed to perform QTL mapping in six/eight environments.ResultsA total of 30 QTLs on chromosomes 1A, 1B, 1D, 2A, 2B, 2D, 3A, 4A, 5A, 5B, 6A, 6D, 7A, 7B and 7D were identified. Among them, six major QTLs QTgw.cib-6A.1, QTgw.cib-6A.2, QGw.cib-6A, QGl.cib-3A, QGl.cib-6A, and QSl.cib-2D explaining 5.96-23.75% of the phenotypic variance were detected in multi-environments and showed strong and stable effects on corresponding traits. Three QTL clusters on chromosomes 2D and 6A containing 10 QTLs were also detected, which showed significant pleiotropic effects on multiple traits. Additionally, three Kompetitive Allele Specific PCR (KASP) markers linked with five of these major QTLs were developed. Candidate genes of QTgw.cib-6A.1/QGl.cib-6A and QGl.cib-3A were analyzed based on the spatiotemporal expression patterns, gene annotation, and orthologous search.ConclusionsSix major QTLs for TGW, GL, GW and SL were detected. Three KASP markers linked with five of these major QTLs were developed. These QTLs and KASP markers will be useful for elucidating the genetic architecture of grain yield and developing new wheat varieties with high and stable yield in wheat.

Nitrogen is a vital element for soil fertility and crop productivity. The transformation of nitrogen is directly affected by tillage practices for the disturbing soil. The characteristics of different nitrogen forms under different tillage modes are still unclear. A 3-year cycle tillage experiment was carried out to assess the combination of rotary tillage (RT), deep tillage (DT), and shallow rotary tillage (SRT) on nitrogen transformation and distribution, wheat yield and nitrogen balance in fluvo-aquic soil from Huang-Huai-Hai Plain in China. The results showed the rotation tillage cycle with deep tillage in the first year increased the total nitrogen (TN), and the main nitrogen form content in 0-30 cm compared with continued rotary tillage (RT-RT-RT). Moreover, the nitrate (NO3--N) and ammonium nitrogen (NH4+-N) content were improved in 20-40 cm by deep tillage practice with the highest value as 39.88 mg kg-1 under DT-SRT-RT. The time, tillage, and depth significantly affected the different nitrogen forms, but there was no effect on dissolved organic carbon (DON) and soil microbial biomass nitrogen (SMBN) by the interaction of time and tillage. Moreover, compared with RT-RT-RT, the rotation tillage promoted the spike number and kernels per spike of wheat, further increasing the wheat yield and nitrogen partial productivity, and with a better effect under DT-SRT-RT. The NO3--N and NH4+-N trended closer and positively correlated with wheat yield in 0-40 cm in 2019. The rotation tillage with deep tillage improved the different forms of nitrogen in 0-30 cm, wheat yield, and nitrogen partial productivity, and decreased the apparent nitrogen loss. It was suggested as the efficiency tillage practice to improve nitrogen use efficiency and crop yield in this area.

Over the past century, environmental changes have significantly impacted wheat spike morphology, crucial for adaptation and grain yield. However, the changes in wheat spike modifications during this periodremain largely unknown. This study examines 16 spike morphology traits in 830 accessions released from 1900 to 2020. It finds that spike weight, grain number per spike (GN), and thousand kernel weight have significantly increased, while spike length has no significant change. The increase in fertile spikelets is due to fewer degenerated spikelets, resulting in a higher GN. Genome-wide association studies identified 49,994 significant SNPs, grouped into 293 genomic regions. The accumulation of favorable alleles in these genomic regions indicates the genetic basis for modification in spike morphology traits. Genetic network analysis of these genomic regions reveals the genetic basis for phenotypic correlations among spike morphology traits. The haplotypes of the identified genomic regions display obvious geographical differentiation in global accessions and environmental adaptation over the past 120 years. In summary, we reveal the genetic basis of adaptive evolution and the interactions of spike morphology, offering valuable resources for the genetic improvement of spike morphology to enhance environmental adaptation.

Twenty-eight field experiments on sandy-loam soils in the UK (1982–2003) are reviewed by relating the extension of the green area duration of the flag leaf (GLADF) by fungicides to effects on yield and quality of winter wheat. Over all experiments mean grain yield =8.85 t ha-1 at 85% DM. With regards quality, mean values were: thousand grain weight (TGW)=44.5 g; specific weight (SWT)=76.9 kg hl-1; crude protein concentration (CP (N× 5.7))=12.5% DM; Hagberg falling number (HFN)=285 s; and sodium dodecyl sulphate (SDS)-sedimentation volume=69 ml. For each day (d) that fungicides increased GLADF there were associated average increases in yield (0.144 t ha-1 d-1, se = 0.0049, df=333), TGW (0.56 g d-1, se=0.017) and SWT (0.22 kg hl-1d-1, se=0.011). Some curvature was evident in all these relationships. When GLADF was delayed beyond 700 ˆCd after anthesis, as was possible in cool wet seasons, responses were curtailed, or less reliable. Despite this apparent terminal sink limitation, fungicide effects on sink size, eg endosperm cell numbers or maximum water mass per grain, were not prerequisites for large effects on grain yield, TGW or SWT. Fungicide effects on CP were variable. Although the average response of CP was negative (-0.029% DM/d; se=0.00338), this depended on cultivar and disease controlled. Controlling biotrophs such as rusts (Puccinia spp.) tended to increase CP, whereas controlling a more necrotrophic pathogen (Septoria tritici) usually reduced CP. Irrespective of pathogen controlled, delaying senescence of the flag leaf was associated with increased nitrogen yields in the grain (averaging 2.24 kg N ha-1d-1, se=0.0848) due to both increased N uptake into the above ground crop, and also more efficient remobilisation of N from leaf laminas. When sulphur availability appeared to be adequate, fungicide x cultivar interactions were similar on S as for CP, although N:S ratios tended to decline (i.e. improve for bread making) when S. tritici was controlled. On average, SDS-sedimentation volume declined (-0.18 ml/d, se=0.027) with increased GLADF, broadly commensurate with the average effect on CP. Hagberg falling number decreased as fungicide increased GLADF (-2.73 s/d, se=0.178), indicating an increase in alpha-amylase activity

Improving yield stability under water-limited conditions is a key objective of wheat breeding programmes. One trait of particular interest is carbohydrate accumulation and remobilisation. This study assessed the genetic basis of aspects of yield and flag leaf sugar contents under drought and well-watered conditions using QTL mapping in a population of 90 doubled haploid lines derived from the cross Chinese Spring × SQ1. As well as soluble sugar content, glucose, fructose, sucrose, and maltose, the traits grain yield (Yld), biomass (Bio), and thousand grain weight (TGW) were also analysed. Analysis of variance showed that genotype, environment and their interactions significantly influenced all the traits studied, with environmental effects explaining up to 74.4% of the total variation. QTL analysis identified 40 QTLs for Yld, TGW, and Bio as well as 53 QTLs for soluble carbohydrates, accounting for up to 40% of phenotypic variation. QTLs coincident for more than one trait were identified on 21 chromosome regions, associated with carbohydrate metabolism and yield performance under drought, particularly on chromosomes 2D, 4A, 4B, 5B, 5D, 6B, and 7A. Candidate genes for several yield-related QTLs were identified. These results provide useful genetic markers for the development of more drought-resistant wheat cultivars.

Spike morphology is crucial for wheat (Triticum aestivum L.) yield and environmental adaptation. We developed a high-throughput phenotyping platform to dissect spike morphology traits based on 54 traits in 1,359 wheat accessions. These 54 spike morphology traits exhibited clear geographical differences among 306 worldwide accessions and breeding selection trend across different time windows for 1,053 accessions released from 1900 to 2020 in China. Based on geographical distribution and breeding selection of haplotypes, we attribute the differences in spike morphology to variable haplotype combinations. Wheat breeding breaks the trade-off between spike length and width/thickness, resulting in increased spike volume. A large proportion of genomic regions has been identified across wheat varieties and utilized as a fixed group to facilitate the targeted improvement and selection of desirable traits during wheat breeding programs. Overall, we provide a resource for the molecular design of spike morphology to facilitate future wheat breeding.

Fusarium head blight (FHB) is a major fungal disease that causes severe yield and quality loss in wheat. Biological control can be integrated with other management strategies to control FHB. For this purpose, Trichoderma gamsii strain T6085 is a potential biocontrol agent to limit the infection of F. graminearum and F. culmorum in wheat. However, the possible impacts of T. gamsii T6085 on the broader microbiome associated with the wheat plant are not currently understood. Therefore, we identified bacteria and fungi associated with different wheat tissues, including assessment of their relative abundances and dynamics in response to the application of T6085 and over time, using amplicon sequencing. Residues of the prior year’s wheat crop and the current year’s wheat spikes were collected at multiple time points, and kernel samples were collected at harvest. DNA was extracted from the collected wheat tissues, and amplicon sequencing was performed to profile microbiomes using 16S v4 rRNA amplicons for bacteria and ITS2 amplicons for fungi. Quantitative PCR was performed to evaluate the absolute abundances of F. graminearum and T. gamsii in different wheat tissues. Disease progression was tracked visually during the growing season, revealing that FHB severity and incidence were significantly reduced when T6085 was applied to wheat spikes at anthesis. However, treatment with T6085 did not lessen the F. graminearum abundance in wheat spikes or kernels. There were substantial changes in F. graminearum abundance over time; in crop residue, pathogen abundance was highest at the initial time point and declined over time, while in wheat spikes, pathogen abundance increased significantly over time. The predominant bacterial taxa in wheat spikes and kernels were Pseudomonas, Enterobacter, and Pantoea, while Alternaria and Fusarium were the dominant fungal groups. Although the microbiome structure changed substantially over time, there were no community-scale rearrangements due to the T6085 treatment. The work suggests several other taxa that could be explored as potential biocontrol agents to integrate with T6085 treatment. However, the timing and the type of T6085 application need to be improved to give more advantages for T6085 to colonize and reduce the F. graminearum inoculum in the field.

1. This study used 4 wheat cultivars (Brigadier, Chaucer, Consort, Reaper) from three locations (Crossnacreevy, C; Downpatrick, D; Limavady, L), which had given rise to differences in wheat specific weight (SW), to examine the relationships between apparent metabolisable energy (AME) concentration, broiler performance and wheat SW. 2. The diets contained (g/kg): wheat 744, casein 142, blended vegetable fat 50, dicalcium phosphate 22, potassium bicarbonate 10.8, sodium bicarbonate 7.5, arginine 5, methionine 2, binder 8, trace minerals/vitamins 7.2, titanium dioxide 1.5. The diets were heat-treated (80°C for 2 min) prior to pelleting (3 mm die). 3. SW ranged from 63 to 77 kg/hectolitre (hl), averaging 66, 69 and 76 kg/hl at D, C and L, respectively. In vitro viscosity of the wheat samples ranged from 5.2 to 17.5 cps and thousand grain weight (TGW) from 33.4 to 47.3 g. Mean TGW was similar at C and D (38.7, 37.0 g) but higher at L (43.1 g). In vitro viscosity was similar at C and L (11.2, 10.2 cps) but somewhat higher at D (14.4 cps). Crude protein (6.25 N) ranged from 116 to 147 g/kg and tended to be higher at D. Starch, which ranged from 612 to 656 g/kg, was least at D (617 g/kg) and greatest at L (641 g/kg). 4. Crude protein, crude fibre and total non-starch polysaccharide (NSP) were negatively correlated with SW, the R 2 being respectively 0.38 (P <0.05), 0.16 (NS) and 0.45 (P <0.05). TGW and starch concentration were positively correlated with SW (R 2=0.70, 0.44, respectively). There was a weak (NS) negative relationship (R 2=0.19) between in vitro viscosity and SW. For both TGW and in vitro viscosity, correlations improved when variety was taken into account (R 2=0.95, 0.92, respectively). 5. There were no significant effects of variety on dry matter (DM) intake or live weight gain (LWG). Gain: food was significantly higher (P <0.05) for Consort than for the other three varieties and the metabolisable energy ratio (ME:GE) just failed to attain significance (P =0.062). Calculated wheat AME (MJ/kg DM) was significantly (P <0.05) higher for Consort than for the other three wheats. There was a good correlation (R 2=0.49) for the total data set between gain:food and ME:GE. In vivo viscosity varied from 13.6 to 28.6 cps for individual treatments and was significantly affected by variety (P <0.001). 6. Although there were no significant differences in DM intake or LWG due to site the values for L (SW 76) were 6 and 5% lower, respectively, than for D (SW 66). Gain:food was lower (P <0.05) for C (SW 69) than for D. ME:GE, wheat AME and ME:gain were not significantly different between sites. 7. There was a weak (R 2=0.18) positive relationship between ME:GE and SW corresponding to a 2.5% increase in energy value for a 10 kg/hl increase in SW and no relationship between gain:food and SW. When variety was taken into the regression the slope was similar but R 2 increased to 0.82. 8. ME:GE and wheat AME concentration were negatively correlated with wheat in vitro viscosity (R 2=0.64, 0.55, respectively). 9. It was concluded that in vitro viscosity appears to provide a better basis than SW for prediction of the nutritive value of wheats of unknown variety. If the variety is known then SW could be used to predict energy value. However, the effect of quite a large change in SW (10 kg/hl) was relatively