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Sucrose non-fermenting 1-related protein kinase 2 (SnRK2) is a plant-specific serine/threonine kinase involved in response to adverse environmental stimuli. Previous studies showed that TaSnRK2.4 was involved in response to abiotic stresses and conferred enhanced tolerance to multiple stresses in Arabidopsis. Further experiments were performed to decipher the underlying mechanisms and discover new functions. The genomic sequences of TaSnRK2.4s locating on chromosome 3A, 3B and 3D were obtained. Sequencing identified one and 13 variations of TaSnRK2.4–3A and TaSnRK2.4-3B, respectively, but no variation was detected in TaSnRK2.4-3D. The markers 2.4AM1, 2.4BM1 and 2.4BM2 were developed based on three variations. Association analysis showed that both TaSnRK2.4-3A and TaSnRK2.4-3B were significantly associated with thousand-kernel weight (TKW), and that SNP3A-T and SNP3B-C were favorable alleles for higher TKW. Yeast two-hybrid and split luciferase assays showed that TaSnRK2.4 physically interacted with abiotic stress responsive protein TaLTP3, suggesting that TaSnRK2.4 enhanced abiotic stress tolerance by activating TaLTP3. Our studies suggested that TaSnRK2.4 have potential in improving TKW and response to abiotic stress.

Background Premature senescence of flag leaf severely affects wheat yield and quality. Chlorophyll (Chl) degradation is the most obvious symptom during leaf senescence and catalyzed by a series of enzymes. Pheophytin pheophorbide hydrolase (Pheophytinase, PPH) gene encodes a Chl degradation hydrolase. Results In this study, the coding, genomic and promoter sequences of wheat TaPPH-A gene were cloned. The corresponding lengths were 1467 bp, 4479 bp and 3666 bp, respectively. Sequence structure analysis showed that TaPPH-A contained five exons and four introns. After the multiple sequences alignment of TaPPH-A genome from 36 accessions in a wheat diversity panel, four SNPs and one 2-bp InDel were observed, which formed two haplotypes, TaPPH-7A-1 and TaPPH-7A-2. Based on the SNP at 1299 bp (A/G), a molecular marker TaPPH-7A-dCAPS was developed to distinguish allelic variation (A/G). Using the molecular markers, 13 SSR, and 116 SNP markers, a linkage map of chromosome 7A were integrated. TaPPH-A was mapped on the chromosome region flanked by Xwmc9 (0.94 cM) and AX-95634545 (1.04 cM) on 7A in a DH population. Association analysis between TaPPH-7A allelic variation and agronomic traits found that TaPPH-7A was associated with TGW in 11 of 12 environments and Chl content at grain-filling stage under drought stress using Population 1 consisted of 323 accessions. The accessions possessed TaPPH-7A-1 (A) had higher TGW and Chl content than those possessed TaPPH-7A-2 (G), thus TaPPH-7A-1 (A) was a favorable allelic variation. By analyzing the frequency of favorable allelic variation TaPPH-7A-1 (A) in Population 2 with 157 landraces and Population 3 with 348 modern cultivars, we found it increased from pre-1950 (0) to 1960s (54.5%), then maintained a relatively stable level about 56% from 1960s to 1990s. Conclusion These results suggested the favorable allelic variation TaPPH-7A-1 (A) should be valuable in enhancing grain yield by improving the source (chlorophyll content) and sink (the developing grain) simultaneously. Furthermore, the newly developed molecular marker TaPPH-7A-dCAPS could be integrated into a breeding kit of screening high TGW wheat for marker-assisted selection. Electronic supplementary material The online version of this article (10.1186/s12870-019-1901-0) contains supplementary material, which is available to authorized users.

Fructans are major nonstructural carbohydrates in wheat (Triticum aestivum L.). Fructan 1-fructosyltransferase (1-FFT) is the key enzyme in fructan biosynthesis. In the present study, 96 sequence variants were detected in the 1-FFT-A1 gene among 26 wheat accessions including UR208, and 15 of them result in amino acid substitutions, forming four haplotypes. Two markers M39 and M2164 were developed based on the InDel21-39 and SNP-2164 polymorphisms to distinguish the three haplotypes in the 1-FFT-A1. 1-FFT-A1 was located on chromosome 4A using marker M2164 and was flanked by markers Xcwm27 and 6-SFT-A1. By association analysis using a natural wheat population consisted of 154 accessions, the results showed that the two markers were significantly associated with water-soluble carbohydrate (WSC) content in the lower internode stem and total stem at the early and middle grain filling stages, 1000-grain weight (TGW) at different grain filling stages and peduncle length (PLE). Comparison of the effects of three haplotypes on agronomic traits indicated that TGW, PLE and total number of spikelets per spike (TNSS) were significantly influenced by haplotypes. HapIII showed a significant positive effect on TGW, PLE and TNSS.

Stomatal density and size affect plant water use efficiency, photosynthsis rate and yield. The objective of this study was to gain insights into the variation and genetic basis of stomatal density and size during grain filling under drought stress (DS) and well-watered (WW) conditions. The doubled haploid population derived from a cross of wheat cultivars Hanxuan 10 (H10), a female parent, and Lumai 14 (L14), a male parent, was used for phenotyping at the heading, flowering, and mid- and late grain filling stages along with established amplified fragment length polymorphism (AFLP) and simple sequence repeat (SSR) markers. The stomatal density of doubled haploid (DH) lines was gradually increased, while the stomatal lengths and widths were gradually decreased during grain filling stage. Twenty additive QTLs and 19 pairs of epistatic QTLs for the 3 traits were identified under DS. The other 20 QTLs and 25 pairs epistatic QTLs were obtained under WW. Most QTLs made more than 10% contributions to the total phenotypic variations at one growth stage under DS or WW. Furthermore, QTLs for stomatal density near Xwmc74 and Xgwm291 located on chromosome 5A were tightly linked to previously reported QTLs regulating total number of spikelets per spike, number of sterile spikelets per spike and proportion of fertile spikelets per spike. Qsw-2D-1 was detected across stages, and was in the same marker region as a major QTL for plant height, QPH. cgb-2D.1 . These indicate that these QTLs on chromosomes 5A and 2D are involved in regulating these agronomic traits and are valuable for molecular breeding.

Stress-associated protein (SAP) has functions in maintaining plant cell elongation, embryo development and response to abiotic stresses. TaSAP1-A1 , one of the Triticum aestivum SAP1 ( TaSAP1 ) members located on wheat chromosome 7A was isolated for polymorphism analysis. Hap III of TaSAP1-A1 was found significantly associated with thousand-grain weight (TGW) in multiple environments. In this study, Hap III also made a positive contribution to TGW in Population 2. The distribution of TaSAP1-A1 Hap III was tracked among varieties released in different years and geographical environments of China. The frequency of Hap III showed an increasing trend during the breeding process in two different populations. The Hap III was gradually selected and applied from 6.36% in landraces to 13.50% in modern varieties. These results exhibited that TaSAP1-A1 Hap III was positively selected during wheat breeding, which is beneficial for grain-yield improvement. The preferred Hap III was initially selected and applied in the higher latitude areas of China in accord with the long day season and longer grain filling stage in these areas. Moreover, the frequency of Hap III in recent modern varieties was still quite low (19.29–26.67%). It indicated a high application potential of TaSAP1-A1 Hap III for improving grain yield in wheat breeding.

Drought as a major abiotic stress often occurs from stem elongation to the grain filling stage of wheat in northern China. Plant height (PH) is a suitable trait to model the dissection of drought tolerance. The purposes of the present study were to validate molecular markers for PH developmental behavior and identify elite alleles of molecular markers. After the phenotyping of 154 accessions for PH dynamic development under well-watered (WW) and drought stressed (DS) conditions, and the genotyping of 60 SSR markers from six candidate chromosome regions related to PH found in our previous linkage mapping studies, both parameters PH and drought tolerance coefficient (DTC) calculated by the conditional analysis were used for association mapping. A total of 46 significant association signals (P

Plant cysteine protease (CP) genes are induced by abiotic stresses such as drought, yet their functions remain largely unknown. We isolated the full-length cDNA encoding a Triticum aestivum CP gene, designated TaCP, from wheat by the rapid amplification of cDNA ends (RACE) method. Sequence analysis revealed that TaCP contains an open reading frame encoding a protein of 362 amino acids, which is 96% identical to barley cysteine protease HvSF42. The TaCP transcript level in wheat seedlings was upregulated during polyethylene glycol (PEG) stress, with a peak appearing around 12 h after treatment. TaCP expression level increased rapidly with NaCl treatment at 48 h. TaCP responded strongly to low temperature (4 degree C) treatment from 1 h post-treatment and reached a peak of about 40-fold at 72 h. However, it showed only a very slight response to abscisic acid (ABA). More than one copy of TaCP was present in each of the three genomes of hexaploid wheat and its diploid donors. TaCP fused with green fluorescent protein (GFP) was located in the plasma membrane of onion epidermis cells. Transgenic Arabidopsis plants overexpressing TaCP showed stronger drought tolerance and higher CP activity under water-stressed conditions than wild-type Arabidopsis plants. The results suggest that TaCP plays a role in tolerance to water deficit.

Plant height (PH) reduction has obviously contributed to lodging resistance and yield increase of wheat (Triticum aestivum L.) in the last decades. The objectives of this study were to understand the diversity status of wheat cultivars in northern China and to identify simple sequence repeat (SSR) markers associated with PH. Using 117 SSR markers evenly distributed on wheat chromosomes, a total of 1484 alleles were detected in 136 winter wheat cultivars that were historically planted in northern China. The average allele number per locus ranged from 2 to 42 with an average of 12.6. The polymorphism information content (PIC) values ranged from 0.016 to 0.941 with an average of 0.640. Based on the cluster tree, the cultivars developed by the same breeders or kin cultivars were gathered in the same subgroup. Some cultivars with similar ecotypes or developed contemporaneously also fell into the same subgroup. Nineteen SSRs each were significantly associated with PH under drought and well-watered conditions, respectively (P

Parameters of chlorophyll fluorescence kinetics (PCFKs) under drought stress condition are generally used to characterize instincts for dehydration tolerance in wheat (Triticum aestivum L.). Therefore, it is important to map quantitative trait loci (QTLs) for PCFKs in wheat genetic improvement for drought tolerance. A doubled haploid (DH) population with 150 lines, derived from a cross between two common wheat varieties, Hanxuan 10 and Lumai 14, was used to analyze the correlation between PCFKs and chlorophyll content (ChlC) and to map QTLs at the grain-filling stage under conditions of both rainfed (drought stress, DS) and well-watered (WW), respectively. QTLs for these traits were detected by QTLMapper version 1.0 based on the composite interval mapping method of the mixed-linear model. The results showed a very significant positive correlation between Fv, Fm, Fv/Fm and Fv/Fo. The correlation coefficients were generally higher under WW than under DS. Also, there was a significant or a highly significant positive correlation between Fv, Fm, Fv/Fm, Fv/Fo and ChlC. The correlation coefficients were higher in the DS group than the WW group. A total of 14 additive QTLs (nine QTLs detected under DS and five QTLs under WW) and 25 pairs of epistatic QTLs (15 pairs detected under DS and 10 pairs under WW) for PCFKs were mapped on chromosomes 6A, 7A, 1B, 3B, 4D and 7D. The contributions of additive QTLs for PCFKs to phenotype variation were from 8.40% to 72.72%. Four additive QTLs (two QTLs detected under DS and WW apiece) controlling ChlC were mapped on chromosomes 1A, 5A and 7A. The contributions of these QTLs for ChlC to phenotype variation were from 7.27% to 11.68%. Several QTL clusters were detected on chromosomes 1B, 7A and 7D, but no shared chromosomal regions for them were identified under different water regimes, indicating that these QTLs performed different expression patterns under rainfed and well-watered conditions. (Handling editor: Yong-Biao Xue)

Summary TaTEF-7A is a functional regulatory factor gene for grain number and other traits, and its diagnostic markers could be used in marker-assisted selection to improve yield potential in wheat., In this study, TaTEF-7A, a member of the transcript elongation factor gene family, and its flanking sequences were isolated. TaTEF-7A was located on chromosome 7A and was flanked by markers Xwmc83 and XP3156.3. Subcellular localization revealed that TaTEF-7A protein was localized in the nucleus. This gene was expressed in all organs, but the highest expression occurred in young spikes and developing seeds. Overexpression of TaTEF-7A in Arabidopsis thaliana produced pleiotropic effects on vegetative and reproductive development that enhanced grain length, silique number, and silique length. No diversity was found in the coding region of TaTEF-7A, but 16 single nucleotide polymorphisms and Indels were detected in the promoter regions of different cultivars. Markers based on sequence variations in the promoter regions (InDel-629 and InDel-604) were developed, and three haplotypes were identified based on those markers. Haplotype–trait association analysis of the Chinese wheat mini core collection revealed that TaTEF-7A was significantly associated with grain number per spike. Phenotyping of near-isogenic lines (NILs) confirmed that TaTEF-7A increases potential grain yield and yield-related traits. Frequency changes in favoured haplotypes gradually increased in cultivars released in China from the 1940s. Geographic distributions of favoured haplotypes were characterized in six major wheat production regions worldwide. The presence of Hap-7A-3, the favoured haplotype, showed a positive correlation with yield in a global set of breeding lines. These results suggest that TaTEF-7A is a functional regulatory factor for grain number per spike and provide a basis for marker-assisted selection.

Taking one hundred and fifty doubled haploid (DH) lines of winter wheat cultivars Hanxuan 10 x Lumai 14 as well as their parents as test materials, the genetic bases of top three leaves length, breadth, and angle at mid-filling stage, and their correlations with yield traits were studied under two water regimes rainfed (drought stress, DS) and well-irrigation (WI) in 2005-2007. Under DS, the leaf length and width of DH lines and their parents were significantly lower than those under WI, whereas the leaf angle had a miscellaneous variation. Under the two water regimes, all the test traits of DH lines had a transgressive segregation, with the variation coefficients ranged from 5.1% to 45.9%. The heritability and gene numbers for the given traits showed great differences. Under WI and DS, the heritability of flag leaf angle (FLA) was the highest (91% and 97%, respectively), and that of the third leaf angle (TLA) was the lowest (23% and 31%, respectively). However, the gene number for FLA (4 and 2 under WI and DS, respectively) was the least, and that of TLA (21 and 25, respectively) in 2007 was the most. Interactive effects were observed among the genes controlling FLA, second leaf angle (SLA), and TLA under WI and DS, and controlling third leaf length (TLL) under WI. The grain number per spike and the grain weight per spike were positively correlated with the length and width of top three leaves, and the thousand-grain weight and the yield per plant were significantly correlated with TLL, FLA, and SLA, but all correlation coefficients were smaller (0.481). It was suggested that the selection of top three leaves length and width should be carried out in the early generations of breeding procedure, while that of leaf angles should be carried out in advanced generations. An appropriate soil moisture regime retained in the critical growth period of the leaves would promote their growth and contribute to the grain yield.

Genetic analyses of nine traits associated with stem water-soluble carbohydrate (SWSC) accumulation and remobilization at grain-filling period under drought stress (DS) and well-watered (WW) conditions were undertaken using doubled haploid lines (DHLs) derived from two Chinese common wheat cultivars. Some significantly and very significantly positive correlation was observed among nine traits associated with SWSC. Higher phenotypic values for most traits were detected under DS. Broad sense heritabilities (\documentclass[10pt]{article} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \usepackage{pmc} \pagestyle{empty} \oddsidemargin -1.0in \begin{document} \begin{equation*}h_{{\mathrm{B}}}^{2}\end{equation*}\end{document}) of the traits showed wide fluctuations between two water treatments. A total of 48 additive and 62 pairs of epistatic QTL for nine traits were identified as distributing on all 21 chromosomes. A majority of QTL involved significant additive and epistatic effects with interactions of QTL and environments (QEIs). Two additive and two pairs of epistatic loci involved only QEIs without corresponding significant additive or epistatic effects. The contributions of the additive QEIs were two- to fourfolds higher than those of their corresponding additive QTL. Most of the additive QEIs for traits associated with SWSC interacted with DS. In addition, some QTL for the grain-filling efficiencies and thousand-grain weight were colocated in the same or adjacent chromosome intervals with QTL for accumulation and remobilization efficiency of SWSC before 14 days after flowering.