• <tr id="yyy80"></tr>
  • <sup id="yyy80"></sup>
  • <tfoot id="yyy80"><noscript id="yyy80"></noscript></tfoot>
  • 99热精品在线国产_美女午夜性视频免费_国产精品国产高清国产av_av欧美777_自拍偷自拍亚洲精品老妇_亚洲熟女精品中文字幕_www日本黄色视频网_国产精品野战在线观看 ?

    A wheat chromosome 5AL region confers seedling resistance to both tan spot and Septoria nodorum blotch in two mapping populations

    2019-12-20 06:40:12WenjingHuXinyaoHeSusanneDreisigackerCarolinaSansaloniPhilominJulianaPawanSingh
    The Crop Journal 2019年6期

    Wenjing Hu, Xinyao He,Susanne Dreisigacker,Carolina P.Sansaloni,Philomin Juliana, Pawan K. Singh,*

    aKey Laboratory of Wheat Biology and Genetic Improvement for Low&Middle Yangtze Valley,Ministry of Agriculture,Lixiahe Region Institute of Agricultural Sciences of Jiangsu Province,Yangzhou 225007,Jiangsu,China

    bInternational Maize and Wheat Improvement Center(CIMMYT), Apdo.Postal 6-64a,06600 Mexico D.F., Mexico

    Keywords:Parastagonospora nodorum Pyrenophora tritici-repentis QTL mapping Resistance breeding Triticum aestivum

    ABSTRACT Tan spot (TS) and Septoria nodorum blotch (SNB), caused by Pyrenophora tritici-repentis and Parastagonospora nodorum, respectively, are important fungal leaf-spotting diseases of wheat that cause significant losses in grain yield. In this study, two recombinant inbred line populations, ‘Bartai' × ‘Ciano T79' (referred to as B × C) and ‘Cascabel' × ‘Ciano T79'(C × C)were tested for TS and SNB response in order to determine the genetic basis of seedling resistance. Genotyping was performed with the DArTseq genotypingby-sequencing (GBS) platform. A chromosome region on 5AL conferred resistance to TS and SNB in both populations,but the effects were larger in B × C(R2 = 11.2%-16.8%)than in C × C(R2 = 2.5%-9.7%).Additionally,the chromosome region on 5BL(presumably Tsn1)was significant for both TS and SNB in B × C but not in C × C.Quantitative trait loci(QTL)with minor effects were identified on chromosomes 1B,2A,2B,3A,3B,4D,5A,5B,5D,6B,and 6D. The two CIMMYT breeding lines ‘Bartai' and ‘Cascabel' contributed resistance alleles at both 5AL and 5BL QTL mentioned above. The QTL on 5AL showed linkage with the Vrn-A1 locus, whereas the vrn-A1 allele conferring lateness was associated with resistance to TS and SNB.

    1. Introduction

    Tan spot(TS) caused by Pyrenophora tritici-repentis [anamorph:Drechslera tritici-repentis(Died.)Shoem.]and Septoria nodorum blotch (SNB) caused by Phaeosphaeria nodorum [anamorph:Parastagonospora nodorum (Berk.) Castellani & Germano][1,2]are leaf spotting diseases of common(Triticum aestivum L.)and durum (T. turgidum L. var. durum) wheat. Both diseases are important and can be devastating in many wheat producing regions worldwide[3-7].

    A number of practices have been used to manage these diseases, including rotations with non-host crops and destruction or avoidance of infested straw, stubble, and volunteer plants by either burning or burying[8].Fungicides are also effective, but their use is not cost effective when grain prices are low. In parallel with resistance genes, fungicides exert high selection pressure on pathogen populations, and their excessive or continual use leads to the possibility of fungal resistance/tolerance. Resistant varieties combined with appropriate cultural practices are considered to be the most cost-effective and environmentally benign way to manage these diseases[9].

    Both qualitative and quantitative resistance have been reported for TS and SNB [10]. P. tritici-repentis is known to produce at least three host-specific toxins that interact with corresponding specific host sensitivity genes to cause necrosis and/or extensive chlorosis associated with susceptibility[4].Ptr ToxA causes necrosis and its corresponding sensitivity gene Tsn1 is located on chromosome 5BL[11].Ptr ToxB and Ptr ToxC cause chlorosis, and their sensitivity genes are Tsc2 on chromosome arm 2BS[12]and Tsc1 on chromosome arm 1AS[13],respectively.Quantitative trait loci(QTL)for TS resistance have been mapped on several wheat chromosomes [4].Shankar and co-authors reported that QTL on chromosomes 1A and 2A had major effects on TS resistance and behaved additively to tsn1 on 5BL[14].

    Eight P. nodorum necrotrophic effectors (NEs) or toxins(SnToxA, SnTox1, SnTox2, SnTox3, SnTox4, SnTox5, SnTox6 and SnTox7) and their corresponding host sensitivity loci(Tsn1, Snn1, Snn2, Snn3-B1 and Snn3-D1, Snn4, Snn5, Snn6 and Snn7) have been described for the wheat-P. nodorum pathosystem [15-17], with the latter loci being located on chromosomes 5BL,1BS,2DS,5BS,5DS,1AS,4BL,6AL,and 2DL,respectively [16,18-24]. Additional resistance QTL have been identified on chromosomes 2A,2B, 5A,and 7A [15].

    Both P. tritici-repentis and P. nodorum produce ToxA. The ToxA gene in the former was obtained from the latter via horizontal gene transfer [25]. Tsn1 has been cloned and Tsn1:ToxA is the best-studied interaction in both pathosystems[11].Normally Tsn1 leads to susceptibility to TS and SNB in the presence of races producing SnToxA,but is not active in some genetic backgrounds, suggesting that race-nonspecific resistance QTL act upstream of the ToxA-Tsn1 interaction,diminishing the effects of Tsn1 [26]. This hypothesis was verified by Kariyawasam et al. [27], who identified a major QTL on 3BL that had an inhibitory epistatic effect on the Ptr ToxA-Tsn1 interaction.

    The objectives of the present study were to 1) map QTL underlying TS and SNB resistance at the seedling stage in two bi-parental populations, and 2) evaluate the effects of a 5AL chromosome region on the response to both diseases, given its linkage with the Vrn-A1 locus.

    2. Materials and methods

    2.1. Plant materials

    Two recombinant inbred line (RIL) populations were used in the present study;the female parents were CIMMYT breeding lines Bartai (pedigree BABAX/LR42//BABAX/3/ERA F2000) and Cascabel (SOKOLL//W15.92/WBLL1), and the shared male parent was Ciano T79 (BUCKY/(SIB)MAYA-74/4/BLUEBIRD//HD-832.5.5/OLESEN/3/CIANO-67/PENJAMO-62). All three parents had a spring growth habit. The ‘Bartai' × ‘Ciano T79'population (hereafter referred to as B × C) comprised 231 F2:7lines, and the ‘Cascabel' × ‘Ciano T79' population (C × C) had 226 F2:7lines. The two populations have been used for mapping spot blotch resistance QTL in our previous studies[28,29].

    2.2. Disease screening

    Mexican P. tritici-repentis (Ptr) isolate MexPtr1 and P. nodorum isolate MexSn4 were used to screen for TS and SNB,respectively. Both isolates are ToxA producers based on inoculation experiments using differential genotypes, infiltration experiments, and PCR with the ToxA genetic marker(data not shown). The isolates were grown on V8-PDA media[30], and conidiospore concentrations for inoculation were adjusted to 4 × 103spores mL-1(MexPtr1) and 1 × 107spores mL-1(MexSn4)[31,32].

    Evaluations of TS and SNB response were conducted on seedlings in a greenhouse at 22 °C day and 18 °C night temperatures with a 16-h photoperiod. Experiments were arranged in a randomized complete block design with two replicates, with four plants per entry grown in plastic containers as experimental units to derive mean values for subsequent analysis. Two trials were carried out for each disease in autumn 2016. Cultivars Erik and Glenlea were included in each trial as resistant and susceptible controls,respectively. Inoculations were performed 14 days after planting when the second leaf was fully expanded. A hand sprayer was used to apply the inocula to seedlings until runoff(about 0.5 mL inoculum per plant).When the leaves were dry,the trays were moved to a humid chamber(RH 100%,20 °C)to facilitate infection. After 24 h, the plants were transferred back to the greenhouse bench. A linear scale of 1-5 was used to evaluate both diseases at seven days post-inoculation(dpi)[30,33,34].

    2.3. Genotyping

    The two populations were genotyped with the DArTseq genotyping-by-sequencing(GBS)platform at Genetic Analysis Service for Agriculture (SAGA) in Guadalajara, Mexico. This genotyping platform is based on a combination of complexity reduction methods developed for array-based DArT and sequencing of resulting representations on next-generation sequencing platforms [35]. Additionally, several KASP markers for genes such as Rht-B1, Rht-D1, Vrn-A1 (Table S1)were applied following Dreisigacker et al. [36]. STS marker fcp623 was used for genotyping the Tsn1 locus following the protocol in Faris et al. [11]. Markers with missing data points>20% and segregation ratios beyond the 0.5-2.0 range were excluded from further analysis. Redundant markers were deleted using the BIN functionality of the ICIMapping ver. 4.1 software[37].

    2.4. Linkage and QTL analysis

    The JoinMap v4.1 software [38]was applied to construct linkage groups (LGs) using the Haldane algorithm with LOD values from 3 to 20 for grouping and the Maximum Likelihood algorithm for ordering within each LG. Chromosome anchoring of LGs was done through blasting GBS marker sequences against the IWGSC Chinese Spring reference genome (v1.0).MapQTL v6.0 software[39]was used for QTL mapping.Interval mapping (IM) was first performed to detect potential QTL for each trait, and then multiple QTL mapping (MQM) for each QTL was executed with the closest linked markers as cofactors. QTL were taken as significant and reported if they exceeded a LOD threshold of 3 in at least one experiment or a threshold of 2 in both experiments. The software MapChart ver. 2.3 [40]was used to draw genetic and physical maps in this study.

    2.5. Statistical analysis

    Analysis of variance (ANOVA) was performed with the AOV functionality in ICIMapping ver.4.1 software[37].Broad-sense heritability estimates were calculated for ANOVA results using the formula h2=), whereis the genetic variance,is the genotype-by-experiment interaction,is the error variance, y is the number of experiments,and r is the number of replicates.

    2.6. Gene survey in the 5AL QTL region

    The JBrowse tool in T3/Wheat (https://triticeaetoolbox.org/wheat/) was used to retrieve information of high confidence genes from the 5AL QTL region, and the corresponding annotation information was obtained from the URGI website(https://urgi.versailles.inra.fr/download/iwgsc/IWGSC_RefSeq_Annotations/v1.0/). Physical positions of markers in IWGSC RefSeq v1.0 were obtained from either T3/Wheat or URGI via BLAST searches.

    3. Results

    3.1. Phenotypic analysis

    Based on ANOVA (Table 1), both diseases showed significant effects of genotype and genotype by environment interaction in the two populations (P <0.001). Broad and continuous variation was observed for both diseases in the two populations (Fig. 1). Although all parental lines showed relatively high resistance, significant variation within the populations was observed, implying different resistance alleles in the parents. Both populations exhibited more resistance to SNB than to TS,and SNB showed higher heritability estimates than TS(Table 1).

    3.2. Genotyping and linkage analysis

    Around 18,000 GBS markers were scored for each population.Additionally, 11 KASP markers were assayed in the B × C population and 13 in the C × C population(Table S1).Tsn1 was polymorphic in the B × C population but monomorphic in the C × C population. After filtering, 1475 and 1798 nonredundant, high quality markers were selected for the B × C and the C × C population, respectively. Using the markers, 35 and 43 linkage groups were generated for the B × C and theC × C population, respectively, representing all 21 wheat chromosomes. The linkage map of B × C covered 4094 cM,with an average marker density of 2.8 cM/marker, whereas the linkage map of C × C covered 5231 cM, with a marker density of 2.9 cM/marker (Table S2).

    Table 1-Analysis of variance for greenhouse experiments of tan spot (TS) and Septoria nodorum blotch (SNB), and their heritability estimates in the‘Bartai' × ‘Ciano T79'(B × C)and ‘Cascabel' × ‘Ciano T79'(C × C)populations.

    Fig.1-Histograms for tan spot(a and c)and Septoria nodorum blotch(b and d)scores in the‘Bartai' × ‘Ciano T79'population(a and b)and the‘Cascabel' × ‘Ciano T79'population(c and d).Disease scores for the resistant check Erick and susceptible check Glenlea are presented,as well as those for the parents,where B stands for‘Bartai',CA for ‘Cascabel',and C for‘Ciano T79'.

    Fig.2- Alignment of the 5AL QTL regions in the ‘Bartai' × ‘Ciano T79'(B × C)and the‘Cascabel' × ‘Ciano T79'(C × C)populations.QTL regions for tan spot(TS)and Septoria nodorum blotch(SNB)and GBS markers are indicated on the outer sides of the linkage groups,whereas marker positions are indicated on the inner sides.Shared markers between the two populations are connected with dotted lines.Linkage groups are only partially presented to show the QTL regions.

    3.3. QTL mapping

    3.3.1. The B × C population

    Two major QTL for TS resistance were detected in the B × C population; one was located on chromosome 5AL in the Vrn-A1 region(Fig.2)and the other on 5BL linking to Tsn1(Fig.S1),accounting on average for 14.2% and 20.3% of phenotypic variation,respectively(Table 2).Resistance alleles at both loci were contributed by the resistant parent Bartai. Additionally,QTL with minor effects were found on chromosomes 3B, 4D and 5B,with resistance alleles from Ciano T79.

    Regarding SNB resistance, the 5AL QTL remained significant, with an average phenotypic variation explained of 16.8%; but the 5BL QTL was less significant, showing an average value of 4.2%.Additional minor QTL were detected on 2B and 4D, with the resistance source being Ciano T79 (Table 2, Fig.S1).

    The 5AL and 5BL QTL act in an additive mode for both diseases (Fig. 3), and the same mode of action was found when all QTL were considered(Fig.S2).

    3.3.2. The C × C population

    Unlike B × C, where major QTL with phenotypic effects >10%were detected, all identified QTL showed minor effects in C × C.A QTL on 5AL was found to be effective against both TS and SNB(Table 3 and Fig.2).Other minor QTL detected in this population included those on 3A,5A,5B and 5D for TS,and 1B,2A,2B,6B and 6D for SNB(Table 3,Fig.S3),none of which was shared between the two diseases nor between the two populations. QTL identified in this population were also in additive in effect(Fig. S2).

    The chromosomal region on 5AL harboring Vrn-A1 was associated with resistance to both diseases in both populations. In B × C, the QTL range for TS and SNB overlapped at Vrn-A1, whereas in C × C, the QTL range for TS was located within that for SNB (Fig. 2). When the projected positions of the markers in the IWGSC Chinese Spring reference genome(v1.0)were used,the QTL for TS and SNB overlapped in B × C,in similar manner to those in C × C,and there was a distance of 2.2 Mb between the two QTL(Fig.4).

    3.4. Gene analysis in the 5AL QTL region

    The 5AL QTL region was defined based on QTL information from the current and a few previous studies,as shown in Fig.4.Physical positions of GBS markers 2341646 and 1204040 that encompassed the core QTL region were used to delimit a region for a survey of annotated genes. The region extending from 582.6 to 589.3 Mb on chromosome 5AL had 88 high confidence genes,including Vrn-A1(Table S4).NBS-LRR genes that are typically associated with disease resistance were not found in the region; however, several genes or gene families that have been reported to be associated with disease resistance were found,including an ABC transporter,Dirigent protein, Receptor-like kinase, Beta-glucosidase, Chaperone DnaJ-domain containing protein, Cysteine protease, F-box family protein, Glucan endo-1,3-beta-glucosidase, Glycosyltransferase, and Pentatricopeptide repeat-containing protein(Table S4), serving as potential candidate genes for this QTL.

    4. Discussion

    In the present study, the 5AL region was significant in both populations for both diseases. Previous studies also reported several QTL on 5AL, mostly for TS. QTs.fcu-5AL was reported by Chu et al. [41]on chromosome 5A between markers barc1061 and cfa2185, explaining up to 18% of variation in seedling experiments.Later,QTs.fcu-5A.1 between barc360 and gwm6 [42]and QYls.lrc-5A closely linked to gdm132 [43]were identified for TS resistance.Recently,QTs.zhl-5A was mapped between markers iwa7025 and iwa5173, accounting for phenotypic variation from 6%to 14%depending on the Ptr isolates assayed [27]. Position comparison of the above-mentioned QTL and those in the current study clearly indicated that the current QTL overlapped with those in Chu et al. [41]andKariyawasam et al.[27](Fig.4).The 5AL QTL regions in Chu et al.[42]and Zwart et al.[43]were extended far from the regions shown in Fig.4 and thus were not drawn;however,their peak regions fall into the same region, as shown in Fig. 4 based on shared markers [42,43]. The Ptr races for which the 5AL QTL showed significant effects were races 1, 2 and 5 in Chu et al.[41], races 1 and 2 in Chu et al. [42], races 1, 2, 3 and 5 in Kariyawasam et al. [27], and race 1 in the current study. It might be concluded that this chromosome region confers a broad spectrum of resistance against Ptr isolates, or that the underlying gene is involved in a yet unknown host-toxin interaction.

    Table 2-QTL for tan spot(TS) and Septoria nodorum blotch(SNB)resistance in the‘Bartai'× ‘Ciano T79'population.

    Fig.3-Phenotypic effects of different combinations of the 5AL and 5BL QTL for TS(a)and SNB(b)in the‘Bartai' × ‘Ciano T79'population.‘+'and‘-' denote the resistant and susceptible alleles,respectively.

    As for SNB,QSnb.fcu-5AL was mapped between barc151 and fcp13, with phenotypic effects of 11% for seedling resistance and 9 to 18% for adult plant resistance [44]. Of its flanking markers,only barc151 was mapped in Fig.4 and thus the distal region could not be determined;it is likely that this QTL region overlapped with the 5AL QTL regions for TS. Additionally,QSnn.niab-5A.1 was found by Cockram et al.[2]and confirmed by Jighly et al. [45], however, its physical location was from 679.5 Mb to 702.5 Mb,beyond the QTL region discussed above.It should also be noted that it was identified in an infiltration experiment using SnTox1, providing further evidence that QSnb.fcu-5AL may be different from QSnn.niab-5A.1.

    The 5AL region has been related to either TS or SNB in previous studies, whereas the current study demonstrated that QTL regions for TS and SNB overlapped in both populations.However,it is still inconclusive if the QTL regions in B × C and C × C represent the same underlying gene. Most likely they are different,due to their positions on the physical map, which are 2.2 Mb apart; but we cannot exclude the possibility that QTL were assigned to a wrong marker interval in one of the populations due to mapping errors, causing the same QTL to appear in different locations in the two populations. As for the possible underlying gene(s) for the QTL, NBS-LRR genes might not be involved considering the absence of such genes in the projected region of the QTL in the Chinese Spring reference genome, although we cannotexclude the possibility that Chinese Spring does not have the resistance gene(s)at all.NBS-LRR genes are generally regarded to be responsible for race-specific resistance against biotrophic diseases [46]; thus it is possible that no such gene was present in the QTL region, considering that TS and SNB are both necrotrophic diseases. Nevertheless, Tsn1, the first identified gene for susceptibility to both TS and SNB,contains an NBS-LRR domain[11],which must be ascribed to its ability to interact specifically with ToxA in both P.tritici-repentis and P.nodorum.In this regard,the underlying gene(s)for the 5AL QTL might not be an NBS-LRR type considering its ability to confer resistance to a wide range of P. tritic-repentis isolates that produce host-specific toxins,as discussed above.

    Table 3-QTL for tan spot(TS)and Septoria nodorum blotch(SNB)resistance in the‘Cascabel'בCiano T79'population.

    Fig.4-Physical locations of markers and QTL regions for tan spot(TS)and Septoria nodorum blotch(SNB)resistance that were reported in the current and previous studies.The physical locations on 5AL were from the IWGSC Chinese Spring RefSeq(v1.0)and shown on the right of the map.QTL confidence intervals were from the respective studies;but the distal region of the SNB QTL in Friesen et al.[44]was unknown,due to the failure in mapping the distal marker cfp13.Vrn-A1 and GBS marker 1141498 that fell in the peak regions for TS and SNB in the present study are in bold and highlighted in red,as well as gdm132 and iwa454 for TS QTL in Chu et al. [41],Zwart et al. [43]and Kariyawasam et al. [30].

    Apart from Tsn1,only two genes have been cloned for leafspotting diseases in wheat, Snn1 for SNB [47]and Stb6 for Septoria tritici blotch (STB) [48], both encoding members of the wall-associated receptor-like kinase family.In this regard,a receptor-like kinase gene was found in the 5AL QTL region and could be a candidate gene. Among other possible candidate genes, ATP-binding cassette (ABC) transporter is famous due to its role in Lr34/Yr18/Sr57/Pm38/Sb1 conferring broad-spectrum resistance in wheat, via transporting or sequestering xenobiotic compounds [49]. In the 5AL QTL region, there are two genes for “ABC transporter B family protein” clustered together at 588.7 Mb, being 1.3 Mb away from Vrn-A1. Dirigent proteins are involved in lignan and lignin biosynthesis as well as in resistance to abiotic and biotic stresses [50], and here we found three dirigent protein genes clustered at 583.6 Mb on 5AL.Interestingly,Juliana et al.[51]also found a TS resistance QTL associated with a dirigent protein, but that one was located on 2AL. Additionally, there are other disease resistance-related genes in the 5AL region,like glucan endo-1,3-beta-glucosidase, glycosyltransferase, F-box family protein, pentatricopeptide repeat-containing protein, cysteine protease and chaperone DnaJ-domain containing protein that are highlighted in Table S4. Nevertheless,further research is needed in order to narrow down the QTL region and number of candidate genes.

    Another new finding of the current study was the localization of vernalization gene Vrn-A1 in this region. It is possible that the gene is associated with disease resistance in the field because Vrn-A1 could contribute to disease escape by its effect on flowering date,as has been reported in Fusarium head blight[52],spot blotch[28]and STB[53].The association of Vrn-A1 with seedling resistance to TS and SNB may imply its linkage with unknown resistance gene(s), although the possibility that Vrn-A1 has pleiotropic effects on disease resistance cannot be ruled out. Limited by the resolution of an ordinary QTL mapping study like the current one, it is difficult to determine the relative positions of the QTL for TS and SNB, and Vrn-A1, for which a fine mapping study on the chromosomal region will be needed.

    The two mapping populations used in the current study have also been characterized for resistance to SB,in which the resistance allele of the 5AL QTL was associated with the vrn-A1 allele that confers lateness[28,29].The same applied to the resistance to TS and SNB in the present study, as well as resistance to FHB and STB as mentioned above, partly explaining why CIMMYT materials generally show resistance to those diseases,at least in Mexican environments,since the vrn-A1 allele is fixed in almost all CIMMYT spring wheat materials [36]. This implies that the utilization of the vrn-A1 allele in breeding might be beneficial in terms of mitigating multiple fungal diseases,as long as the breeding lines are not too late, and this can be achieved by selecting early vrn-A1 lines since there are other genes contributing to earliness.

    Tsn1 has long been recognized as a major factor in experiments where ToxA-Tsn1 interaction happens [11]. In the B × C population, the underlying gene for the 5BL QTL must be Tsn1, considering that both MexPtr1 and MexSn4 produce ToxA,although Tsn1 was not mapped under the peak of the QTL (Fig. S1), which must be ascribed to errors in genotyping of the marker fcp623 that made Tsn1 appear a bit away from its real position. In the C × C population, both parents had the Tsn1 allele for susceptibility, thus explaining why the 5BL QTL was not detected.

    It is noteworthy that unknown host-toxin interaction or unidentified QTL might exist in the two populations analyzed in the current study. It was unexpected that Ciano T79, the Tsn1 carrier, showed moderately resistant reactions to both diseases in both populations. Lines with Tsn1 usually exhibit susceptible reactions when ToxA-producing isolates are inoculated, but some loci may have epistatic effects on Tsn1,at least in the TS system known so far,minimizing the effects of the ToxA-Tsn1 interaction, for example the 3BL QTL reported by Kariyawasam et al. [27]. Another possibility could be that the isolates used in this study secrete unknown toxins that interact with QTL that were not identified in this study. Further research is required to clarify the confounding issues.

    Declaration of Competing Interest

    None.

    Acknowledgments

    This work was supported by the Bill and Melinda Gates Foundation and the United States Agency for International Development (USAID) through the Cereal Systems Initiative for South Asia (CSISA), Durable Rust Resistance in Wheat(DRRW)/Delivering Genetic Gains in Wheat (DGGW) and the CGIAR Research Program for Wheat (CRP WHEAT) project.Technical support from Nerida Lozano was highly appreciated. The first author is grateful for the financial support of The National Key Research and Development Program of China on Molecular Design Breeding in Wheat(2016YFD0101802).

    Supplementary data

    Supplementary data for this article can be found online at https://doi.org/10.1016/j.cj.2019.05.004.

    亚洲美女黄色视频免费看| 久久国内精品自在自线图片| 美女国产视频在线观看| 在线观看免费日韩欧美大片 | 日韩伦理黄色片| 蜜臀久久99精品久久宅男| 欧美xxⅹ黑人| 免费高清在线观看日韩| 黄色一级大片看看| 国产欧美亚洲国产| 一本一本久久a久久精品综合妖精| 国产日韩欧美亚洲二区| 欧美日韩国产mv在线观看视频| 久久精品久久久久久噜噜老黄| 男女边摸边吃奶| 久久免费观看电影| 日韩一本色道免费dvd| 大型av网站在线播放| 一级毛片黄色毛片免费观看视频| 亚洲精品美女久久久久99蜜臀 | 18禁黄网站禁片午夜丰满| 在线观看免费午夜福利视频| 亚洲人成电影观看| 波多野结衣一区麻豆| 亚洲美女黄色视频免费看| 日韩av不卡免费在线播放| 国产成人精品久久二区二区91| 亚洲欧美精品自产自拍| 日韩熟女老妇一区二区性免费视频| 亚洲人成网站在线观看播放| 国产成人系列免费观看| 狠狠精品人妻久久久久久综合| 在线天堂中文资源库| 亚洲精品成人av观看孕妇| 国产免费一区二区三区四区乱码| av在线播放精品| 夫妻午夜视频| 99久久精品国产亚洲精品| 一级毛片 在线播放| 精品国产乱码久久久久久小说| 欧美日韩福利视频一区二区| 久久免费观看电影| 又大又爽又粗| 男人操女人黄网站| 亚洲天堂av无毛| 亚洲五月婷婷丁香| 精品一区二区三区四区五区乱码 | 久久影院123| 99国产精品99久久久久| 亚洲欧美清纯卡通| 下体分泌物呈黄色| 久久国产精品影院| 啦啦啦啦在线视频资源| 久久精品久久久久久久性| 天天躁夜夜躁狠狠躁躁| 搡老岳熟女国产| 午夜免费观看性视频| 久久久久精品人妻al黑| 国产欧美日韩精品亚洲av| 亚洲人成77777在线视频| 国产欧美日韩精品亚洲av| 在线精品无人区一区二区三| 高清视频免费观看一区二区| 日本黄色日本黄色录像| www.999成人在线观看| 国产亚洲欧美在线一区二区| 国产av精品麻豆| 欧美精品人与动牲交sv欧美| 91国产中文字幕| 免费在线观看黄色视频的| 亚洲欧美成人综合另类久久久| 亚洲伊人久久精品综合| 国产一区二区在线观看av| 国产亚洲欧美精品永久| 精品一区二区三区av网在线观看 | 美女中出高潮动态图| 精品人妻在线不人妻| 青春草视频在线免费观看| 十八禁网站网址无遮挡| 国产精品av久久久久免费| 天天躁夜夜躁狠狠久久av| 视频在线观看一区二区三区| 欧美日韩视频精品一区| 首页视频小说图片口味搜索 | 999久久久国产精品视频| 亚洲国产欧美日韩在线播放| 日本91视频免费播放| 国产免费又黄又爽又色| 80岁老熟妇乱子伦牲交| 18禁国产床啪视频网站| 国产欧美日韩精品亚洲av| 亚洲一码二码三码区别大吗| 视频在线观看一区二区三区| 人人妻人人澡人人看| 又粗又硬又长又爽又黄的视频| 国产高清videossex| 黄色a级毛片大全视频| 天天躁日日躁夜夜躁夜夜| 国产在视频线精品| 一二三四在线观看免费中文在| 黑人巨大精品欧美一区二区蜜桃| 午夜福利乱码中文字幕| 日本黄色日本黄色录像| 天天操日日干夜夜撸| 香蕉国产在线看| 十分钟在线观看高清视频www| 一级片'在线观看视频| 久久青草综合色| 91九色精品人成在线观看| 午夜激情av网站| 黑人巨大精品欧美一区二区蜜桃| 日韩 欧美 亚洲 中文字幕| 亚洲国产日韩一区二区| 国产免费福利视频在线观看| 麻豆国产av国片精品| 日韩大片免费观看网站| 国产精品 欧美亚洲| netflix在线观看网站| 久久久久国产一级毛片高清牌| 汤姆久久久久久久影院中文字幕| 一级片免费观看大全| 亚洲伊人久久精品综合| 丰满迷人的少妇在线观看| 99九九在线精品视频| 国产亚洲av高清不卡| 午夜视频精品福利| 无遮挡黄片免费观看| 人人妻人人爽人人添夜夜欢视频| 精品国产乱码久久久久久男人| 亚洲av国产av综合av卡| 观看av在线不卡| 日韩av免费高清视频| 校园人妻丝袜中文字幕| 69精品国产乱码久久久| 中文字幕人妻熟女乱码| 在线av久久热| 天堂8中文在线网| 一本久久精品| 9热在线视频观看99| 老汉色∧v一级毛片| 欧美在线黄色| 在线天堂中文资源库| 午夜福利在线免费观看网站| 高清黄色对白视频在线免费看| 国产色视频综合| 爱豆传媒免费全集在线观看| 制服人妻中文乱码| 久久精品国产综合久久久| 亚洲成人免费电影在线观看 | 色婷婷av一区二区三区视频| 丰满迷人的少妇在线观看| 无限看片的www在线观看| 一边摸一边做爽爽视频免费| 无遮挡黄片免费观看| 99国产精品免费福利视频| 天堂中文最新版在线下载| 99精品久久久久人妻精品| 大码成人一级视频| 精品卡一卡二卡四卡免费| 亚洲欧美日韩另类电影网站| 天天躁夜夜躁狠狠躁躁| 午夜福利,免费看| 大话2 男鬼变身卡| 一区二区av电影网| 国产精品一区二区免费欧美 | 下体分泌物呈黄色| 亚洲少妇的诱惑av| 欧美激情 高清一区二区三区| 精品国产乱码久久久久久男人| 国产一区二区三区av在线| 国产男女内射视频| 在线观看免费高清a一片| 超碰97精品在线观看| 亚洲伊人色综图| 久久av网站| 亚洲一码二码三码区别大吗| 极品人妻少妇av视频| 黄色a级毛片大全视频| 久热爱精品视频在线9| 亚洲专区中文字幕在线| 深夜精品福利| 欧美精品一区二区大全| 国产精品秋霞免费鲁丝片| 成人18禁高潮啪啪吃奶动态图| 国产老妇伦熟女老妇高清| 国产日韩欧美亚洲二区| 国产在线免费精品| 国产色视频综合| 伦理电影免费视频| 黄色视频在线播放观看不卡| 黑人巨大精品欧美一区二区蜜桃| 亚洲精品美女久久久久99蜜臀 | 曰老女人黄片| 国产成人系列免费观看| 亚洲成人国产一区在线观看 | 又黄又粗又硬又大视频| 一边摸一边做爽爽视频免费| 天天影视国产精品| videosex国产| 男人操女人黄网站| 国产99久久九九免费精品| 丰满饥渴人妻一区二区三| 美女高潮到喷水免费观看| 美女主播在线视频| 国产成人免费无遮挡视频| 午夜精品国产一区二区电影| 黄色视频不卡| 日本午夜av视频| 色综合欧美亚洲国产小说| 亚洲国产最新在线播放| 新久久久久国产一级毛片| 国产精品.久久久| 久久99精品国语久久久| 日韩av免费高清视频| 国产免费视频播放在线视频| 亚洲精品成人av观看孕妇| 精品亚洲成a人片在线观看| 99re6热这里在线精品视频| 母亲3免费完整高清在线观看| 爱豆传媒免费全集在线观看| 最近手机中文字幕大全| 久久久久久久大尺度免费视频| 欧美老熟妇乱子伦牲交| 亚洲精品自拍成人| 午夜久久久在线观看| 日本欧美国产在线视频| av在线老鸭窝| 色网站视频免费| 色婷婷av一区二区三区视频| av国产久精品久网站免费入址| 午夜视频精品福利| 久久久久精品人妻al黑| 欧美日韩视频精品一区| √禁漫天堂资源中文www| 好男人视频免费观看在线| 久久久久久人人人人人| 宅男免费午夜| 电影成人av| 亚洲精品国产一区二区精华液| 天天躁日日躁夜夜躁夜夜| 日韩制服丝袜自拍偷拍| 老汉色av国产亚洲站长工具| 日韩电影二区| 波野结衣二区三区在线| e午夜精品久久久久久久| 高清不卡的av网站| 欧美日韩亚洲高清精品| www.999成人在线观看| 人妻一区二区av| 精品人妻在线不人妻| 大型av网站在线播放| 午夜久久久在线观看| 久久久久网色| 丝袜喷水一区| www.av在线官网国产| 国产伦人伦偷精品视频| 国产黄色免费在线视频| 欧美日韩视频精品一区| 久久久精品国产亚洲av高清涩受| 色婷婷av一区二区三区视频| 日本vs欧美在线观看视频| 亚洲欧洲国产日韩| 国产精品三级大全| 18禁裸乳无遮挡动漫免费视频| 自拍欧美九色日韩亚洲蝌蚪91| av在线播放精品| 亚洲av片天天在线观看| 国产日韩欧美视频二区| 两个人免费观看高清视频| 永久免费av网站大全| 久久久久国产一级毛片高清牌| 99久久综合免费| 亚洲 国产 在线| 国产在线视频一区二区| 亚洲一卡2卡3卡4卡5卡精品中文| 久久影院123| 亚洲欧美日韩另类电影网站| 免费看不卡的av| 国产男女超爽视频在线观看| 久久精品久久久久久噜噜老黄| av在线播放精品| 婷婷成人精品国产| 人妻一区二区av| 欧美日韩成人在线一区二区| 青春草亚洲视频在线观看| 一边摸一边做爽爽视频免费| 国产精品久久久人人做人人爽| 天天操日日干夜夜撸| 国产97色在线日韩免费| 美国免费a级毛片| 丝袜喷水一区| 99国产精品一区二区蜜桃av | 国产激情久久老熟女| a级毛片黄视频| 亚洲少妇的诱惑av| 国产精品av久久久久免费| 成人国语在线视频| 中文字幕精品免费在线观看视频| 精品亚洲成国产av| 亚洲av电影在线进入| bbb黄色大片| 精品国产国语对白av| 日本五十路高清| 日韩熟女老妇一区二区性免费视频| 美女主播在线视频| 免费一级毛片在线播放高清视频 | 国产精品国产av在线观看| 日韩大码丰满熟妇| 日本猛色少妇xxxxx猛交久久| 国产精品一区二区精品视频观看| 国产淫语在线视频| 永久免费av网站大全| 丰满人妻熟妇乱又伦精品不卡| 午夜激情av网站| 久久精品aⅴ一区二区三区四区| 男女边摸边吃奶| 国产精品一区二区在线不卡| 啦啦啦啦在线视频资源| 视频区欧美日本亚洲| 一本色道久久久久久精品综合| 午夜福利影视在线免费观看| 欧美精品高潮呻吟av久久| 香蕉丝袜av| 国产激情久久老熟女| 国产日韩欧美视频二区| 老汉色av国产亚洲站长工具| 国产国语露脸激情在线看| 成人影院久久| 欧美精品啪啪一区二区三区 | 亚洲国产精品成人久久小说| 精品人妻在线不人妻| 1024视频免费在线观看| 成人18禁高潮啪啪吃奶动态图| 999精品在线视频| 欧美日本中文国产一区发布| 极品人妻少妇av视频| 中文字幕精品免费在线观看视频| 国产一区有黄有色的免费视频| 免费观看av网站的网址| 欧美日韩亚洲国产一区二区在线观看 | 精品一品国产午夜福利视频| 视频在线观看一区二区三区| xxxhd国产人妻xxx| 天天躁夜夜躁狠狠躁躁| 你懂的网址亚洲精品在线观看| 国产激情久久老熟女| 蜜桃国产av成人99| 成在线人永久免费视频| 久久久国产一区二区| 国产精品一区二区精品视频观看| 尾随美女入室| bbb黄色大片| 国产精品熟女久久久久浪| 国产精品久久久人人做人人爽| 这个男人来自地球电影免费观看| 久久久久久久精品精品| 新久久久久国产一级毛片| 久久久久久久久久久久大奶| 天堂8中文在线网| www.自偷自拍.com| 黄频高清免费视频| 中文字幕最新亚洲高清| 久久人人97超碰香蕉20202| 最近中文字幕2019免费版| 国产精品免费大片| 日韩中文字幕欧美一区二区 | 国产免费视频播放在线视频| 黄色一级大片看看| 少妇的丰满在线观看| 国产精品免费视频内射| 国产欧美亚洲国产| 国产精品亚洲av一区麻豆| 校园人妻丝袜中文字幕| 人人妻,人人澡人人爽秒播 | 婷婷丁香在线五月| 免费看av在线观看网站| 久久中文字幕一级| 精品一区在线观看国产| 国产欧美日韩一区二区三区在线| 在线看a的网站| 黄色怎么调成土黄色| 999久久久国产精品视频| 人成视频在线观看免费观看| 久久午夜综合久久蜜桃| 韩国精品一区二区三区| 美女脱内裤让男人舔精品视频| 国产一区亚洲一区在线观看| 亚洲成人免费电影在线观看 | 我的亚洲天堂| 日本五十路高清| 1024视频免费在线观看| 国产成人a∨麻豆精品| av欧美777| 日韩伦理黄色片| a级毛片黄视频| 国产一区二区三区综合在线观看| 国产男女内射视频| 99热全是精品| 成在线人永久免费视频| 国产高清国产精品国产三级| 久久久久久人人人人人| 永久免费av网站大全| 精品高清国产在线一区| 美女中出高潮动态图| 日韩制服骚丝袜av| 韩国高清视频一区二区三区| 午夜福利在线免费观看网站| www.熟女人妻精品国产| 久久人人97超碰香蕉20202| 老鸭窝网址在线观看| 91精品国产国语对白视频| 久久国产精品男人的天堂亚洲| av福利片在线| 一边摸一边抽搐一进一出视频| 日韩,欧美,国产一区二区三区| av网站在线播放免费| 欧美+亚洲+日韩+国产| 男的添女的下面高潮视频| 高清不卡的av网站| 午夜福利一区二区在线看| 午夜老司机福利片| 亚洲专区国产一区二区| 亚洲av电影在线进入| 成人18禁高潮啪啪吃奶动态图| 天天影视国产精品| 日本一区二区免费在线视频| 亚洲一区中文字幕在线| 美女高潮到喷水免费观看| 欧美国产精品一级二级三级| www.自偷自拍.com| 国产色视频综合| 中文乱码字字幕精品一区二区三区| 久久精品国产亚洲av涩爱| 大码成人一级视频| 中文字幕制服av| 亚洲人成电影观看| 在现免费观看毛片| 久久久国产精品麻豆| 久久午夜综合久久蜜桃| 国产在线观看jvid| 后天国语完整版免费观看| 久久久久网色| 亚洲成色77777| 9色porny在线观看| 精品一区二区三卡| 国产成人91sexporn| 亚洲精品国产区一区二| 伊人亚洲综合成人网| 人妻人人澡人人爽人人| 久久av网站| 午夜激情久久久久久久| 18禁观看日本| 一级,二级,三级黄色视频| 午夜福利影视在线免费观看| 亚洲男人天堂网一区| 美女扒开内裤让男人捅视频| 老司机靠b影院| 日韩av不卡免费在线播放| 亚洲成av片中文字幕在线观看| 午夜福利视频在线观看免费| 嫁个100分男人电影在线观看 | 人人妻人人爽人人添夜夜欢视频| xxx大片免费视频| 久久久久久免费高清国产稀缺| 成年美女黄网站色视频大全免费| av国产精品久久久久影院| 国产主播在线观看一区二区 | 亚洲精品成人av观看孕妇| 天天添夜夜摸| 亚洲欧美日韩高清在线视频 | 国产成人av激情在线播放| 亚洲欧美色中文字幕在线| 国产国语露脸激情在线看| 丰满饥渴人妻一区二区三| 亚洲欧美精品自产自拍| 人人妻人人添人人爽欧美一区卜| 天天影视国产精品| 久久这里只有精品19| 每晚都被弄得嗷嗷叫到高潮| 国产福利在线免费观看视频| 在线观看免费高清a一片| 免费在线观看完整版高清| 亚洲精品国产一区二区精华液| 国产精品久久久久久精品电影小说| 国产精品.久久久| 精品一区二区三区av网在线观看 | 日韩大码丰满熟妇| 97人妻天天添夜夜摸| 久久中文字幕一级| 久久久久久久精品精品| 91麻豆精品激情在线观看国产 | 国产av精品麻豆| 高潮久久久久久久久久久不卡| 两个人免费观看高清视频| 久久久久国产一级毛片高清牌| 韩国精品一区二区三区| 亚洲熟女精品中文字幕| 麻豆乱淫一区二区| 亚洲精品久久久久久婷婷小说| a 毛片基地| 久久人妻熟女aⅴ| 精品第一国产精品| 国产精品香港三级国产av潘金莲 | 悠悠久久av| 日韩av不卡免费在线播放| 丝袜脚勾引网站| 天堂中文最新版在线下载| 好男人视频免费观看在线| 黄色一级大片看看| 高潮久久久久久久久久久不卡| 在线观看免费视频网站a站| 爱豆传媒免费全集在线观看| 十八禁高潮呻吟视频| 精品国产乱码久久久久久小说| 晚上一个人看的免费电影| 十八禁人妻一区二区| 久久久久国产精品人妻一区二区| 男女下面插进去视频免费观看| 美国免费a级毛片| 亚洲人成77777在线视频| netflix在线观看网站| 国产免费现黄频在线看| 国产av国产精品国产| 看免费成人av毛片| 啦啦啦中文免费视频观看日本| 亚洲国产日韩一区二区| 黄色怎么调成土黄色| 欧美黑人欧美精品刺激| 久久午夜综合久久蜜桃| 亚洲成人国产一区在线观看 | av在线app专区| 日韩制服骚丝袜av| 好男人电影高清在线观看| 后天国语完整版免费观看| 99热全是精品| 国产成人影院久久av| 精品少妇一区二区三区视频日本电影| 大话2 男鬼变身卡| 国产老妇伦熟女老妇高清| 永久免费av网站大全| 亚洲国产精品成人久久小说| cao死你这个sao货| 欧美 亚洲 国产 日韩一| 男女之事视频高清在线观看 | 久久精品国产a三级三级三级| 国产一区二区在线观看av| 人人妻,人人澡人人爽秒播 | 人妻人人澡人人爽人人| 欧美日韩综合久久久久久| 一本久久精品| 在线看a的网站| 曰老女人黄片| 日韩一本色道免费dvd| 久久久久国产精品人妻一区二区| 十八禁人妻一区二区| 欧美日韩黄片免| 久9热在线精品视频| 男人舔女人的私密视频| 日本一区二区免费在线视频| av线在线观看网站| 成人国语在线视频| 男人添女人高潮全过程视频| av网站在线播放免费| 久久久精品免费免费高清| 青春草亚洲视频在线观看| 午夜福利,免费看| 一级毛片女人18水好多 | 久久综合国产亚洲精品| 在线精品无人区一区二区三| 欧美大码av| 亚洲av美国av| 免费一级毛片在线播放高清视频 | 国产男女内射视频| 欧美黑人精品巨大| 青春草亚洲视频在线观看| 亚洲男人天堂网一区| 丁香六月欧美| 亚洲九九香蕉| 亚洲欧美中文字幕日韩二区| 丝袜人妻中文字幕| 国产无遮挡羞羞视频在线观看| 欧美精品一区二区免费开放| 欧美 日韩 精品 国产| 国产成人一区二区三区免费视频网站 | 欧美日韩精品网址| 国产福利在线免费观看视频| 久久久久国产精品人妻一区二区| 成人手机av| 亚洲熟女毛片儿| 精品熟女少妇八av免费久了| 国产亚洲精品第一综合不卡| 午夜激情久久久久久久| 国产精品久久久人人做人人爽| 久久影院123| 亚洲,欧美,日韩| 亚洲中文日韩欧美视频| 免费观看av网站的网址| 久久国产精品影院| 亚洲熟女精品中文字幕| 日韩制服丝袜自拍偷拍| 亚洲成国产人片在线观看| h视频一区二区三区| 国产一区二区 视频在线| www.999成人在线观看| 国产成人免费无遮挡视频| 午夜福利在线免费观看网站| 男男h啪啪无遮挡| 少妇裸体淫交视频免费看高清 | av欧美777| 黄片播放在线免费| 久久女婷五月综合色啪小说| 欧美国产精品va在线观看不卡| 国产免费福利视频在线观看| 少妇 在线观看| 99香蕉大伊视频|