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1.
A major QTL conferring crown rot resistance in barley and its association with plant height 总被引:1,自引:0,他引:1
H. B. Li M. X. Zhou C. J. Liu 《TAG. Theoretical and applied genetics. Theoretische und angewandte Genetik》2009,118(5):903-910
Crown rot (CR) is one of the most destructive diseases of barley and wheat. Fusarium species causing CR survive in crop residue and a growing acceptance of stubble retention practices has exacerbated disease
severity and yield loss. Growing resistant cultivars has long been recognised as the most effective way to reduce CR damage
but these are not available in barley. In a routine screening of germplasm, a barley landrace from China gave the best CR
resistance among the genotypes tested. Using a doubled haploid population derived from this landrace crossed to Franklin,
we demonstrate that the CR resistance of TX9425 was conditioned by a major QTL. The QTL, designated as Qcrs.cpi-3H, was mapped near the centromere on the long arm of chromosome 3H. Its effect is highly significant, accounting for up to
63.3% of the phenotypic variation with a LOD value of 14.8. The location of Qcrs.cpi-3H was coincident with a major QTL conferring plant height (PH) and the effect of PH on CR reaction was also highly significant.
When the effect of PH was accounted for by covariance analysis, the Qcrs.cpi-3H QTL remained highly significant, accounting for over 40% of the phenotypic variation. The existence of such a major QTL implies
that breeding barley cultivars with enhanced CR resistance should be feasible. 相似文献
2.
Identification and fine-mapping of a major QTL conferring resistance against head smut in maize 总被引:4,自引:0,他引:4
Chen Y Chao Q Tan G Zhao J Zhang M Ji Q Xu M 《TAG. Theoretical and applied genetics. Theoretische und angewandte Genetik》2008,117(8):1241-1252
Head smut is one of the most devastating diseases in maize, causing severe yield loss worldwide. Here we report identification
and fine-mapping of a major quantitative trait locus (QTL) conferring resistance to head smut. Two inbred lines ‘Ji1037’ (donor
parent, highly resistant) and ‘Huangzao4’ (recurrent parent, highly susceptible) were crossed and then backcrossed to ‘Huangzao4’
to generate BC populations. Four putative resistance QTLs were detected in the BC1 population, in which the major one, designated as qHSR1, was mapped on bin 2.09. The anchored ESTs, IDPs, RGAs, BAC and BAC-end
sequences in bin 2.09 were exploited to develop markers to saturate the qHSR1 region. The recombinants in the qHSR1 region
were obtained by screening the BC2 population and then backcrossed again to ‘Huangzao4’ to produce 59 BC2:3 families or selfed to generate nine BC2F2 families. Individuals from each BC2:3 or BC2F2 family were evaluated for their resistances to head smut and genotypes at qHSR1. Analysis of genotypes between the resistant
and susceptible groups within the same family allows deduction of phenotype of its parental BC2 recombinant. Based on the 68 BC2 recombinants, the major resistance QTL, qHSR1, was delimited into an interval of ~2 Mb, flanked by the newly developed markers
SSR148152 and STS661. A large-scale survey of BC2:3 and BC2F2 progeny indicated that qHSR1 could exert its genetic effect by reducing the disease incidence by ~25%.
Yongsheng Chen, Qing Chao and Guoqing Tan contributed equally to this work. 相似文献
3.
G. D. Chen Y. X. Liu Y. M. Wei C. L. McIntyre M. X. Zhou Y.-L. Zheng C. J. Liu 《TAG. Theoretical and applied genetics. Theoretische und angewandte Genetik》2013,126(10):2511-2520
Fusarium crown rot (FCR) is a serious cereal disease in semi-arid regions worldwide. In assisting the effort of breeding cultivars with enhanced resistance, we identified several barley genotypes with high levels of FCR resistance. One of these genotypes, AWCS079 which is a barley landrace originating from Japan, was investigated by developing and assessing three populations of recombinant inbred lines. Two QTL, one located on the long arm of chromosome 1H (designated as Qcrs.cpi-1H) and the other on 3HL (designated as Qcrs.cpi-3H), were found to be responsible for the FCR resistance of this genotype. Qcrs.cpi-1H is novel as no other FCR loci have been reported on this chromosome arm. Qcrs.cpi-3H co-located with a reduced height (Rht) locus and the effectiveness of the former was significantly affected by the latter. The total phenotypic variance explained by these two QTL was over 60 %. Significant effects were detected for each of the QTL in each of the three populations assessed. The existence of these loci with major effects should not only facilitate breeding and exploitation of FCR-resistant barley cultivars but also their further characterization based on fine mapping and map-based gene cloning. 相似文献
4.
5.
Micic Z Hahn V Bauer E Melchinger AE Knapp SJ Tang S Schön CC 《TAG. Theoretical and applied genetics. Theoretische und angewandte Genetik》2005,111(2):233-242
Midstalk rot, caused by Sclerotinia sclerotiorum (Lib.) de Bary, is an important cause of yield loss in sunflower (Helianthus annuus L.). Objectives of this study were to: (1) estimate the number, genomic positions and genetic effects of quantitative trait loci (QTL) for resistance to midstalk rot in line TUB-5-3234, derived from an interspecific cross; (2) determine congruency of QTL between this line and other sources of resistance; and (3) make inferences about the efficiency of selective genotyping (SG) in detecting QTL conferring midstalk rot resistance in sunflower. Phenotypic data for three resistance (stem lesion, leaf lesion and speed of fungal growth) and two morphological (leaf length and leaf length with petiole) traits were obtained from 434 F3 families from cross CM625 (susceptible) × TUB-5-3234 (resistant) under artificial infection in field experiments across two environments. The SG was applied by choosing the 60 most resistant and the 60 most susceptible F3 families for stem lesion. For genotyping of the respective F2 plants, 78 simple sequence repeat markers were used. Genotypic variances were highly significant for all traits. Heritabilities and genotypic correlations between resistance traits were moderate to high. Three to four putative QTL were detected for each resistance trait explaining between 40.8% and 72.7% of the genotypic variance (
). Two QTL for stem lesion showed large genetic effects and corroborated earlier findings from the cross NDBLOSsel (resistant) × CM625 (susceptible). Our results suggest that SG can be efficiently used for QTL detection and the analysis of congruency for resistance genes across populations. 相似文献
6.
Genomic regions conferring resistance to multiple fungal pathogens in synthetic hexaploid wheat 总被引:1,自引:0,他引:1
Abdulqader Jighly Manickavelu Alagu Farid Makdis Murari Singh Sukhwinder Singh Livinus C. Emebiri Francis C. Ogbonnaya 《Molecular breeding : new strategies in plant improvement》2016,36(9):127
Fungal diseases are among the most devastating biotic stresses and often cause significant losses in wheat production worldwide. A set of 173 synthetic hexaploid wheat (SHW) characterized for resistance against fungal pathogens that cause leaf, stem and yellow rusts, yellow leaf spot, Septoria nodorum and crown rot were used in genome-wide association study (GWAS). Diversity Arrays Technology (DArT) and DArTSeq markers were employed for marker–trait association in which 74 markers associated with 35 quantitative trait loci (QTL) were found to be significantly linked with disease resistances using a unified mixed model (P = 10?3 to 10?5); Of these 15 QTL originated from D genome. Six markers on 1BL, 3BS, 4BL, 6B, and 6D conferred resistance to two diseases representing 10 of the 35 QTL. A further set of 147 SHW genotyped with DArT only markers validated 11 QTL detected in the previous 173 SHW. We also confirmed the presence of the gene Lr46/Yr29/Sr58/Pm39/Ltn2 on 1BL in the SHW germplasm. In addition, gene–gene interactions between significantly associated loci and all loci across the genome revealed five significant interactions at FDR <0.05. Two significant leaf rust and one stem rust interactions were thought to be synergistic, while another two QTL for yellow leaf spot involved antagonistic relations. To the best of our knowledge, this is the first GWAS for six fungal diseases using SHW. Identification of markers associated with disease resistance to one or more diseases represents an important resource for pyramiding favorable alleles and introducing multiple disease resistance from SHW accessions into current elite wheat cultivars. 相似文献
7.
Haijun Qi Xiuliang Zhu Wenbiao Shen Xia Yang Chaozhong Zhang Genying Li Feng Chen Xuening Wei Zengyan Zhang 《植物学报(英文版)》2024,(1):12-16
<正>Fusarium crown rot (FCR), mainly caused by the soil-borne fungus Fusarium pseudograminearum, is a devastating disease of wheat (Triticum aestivum (Ta)). Fusarium crown rot causes substantial yield losses and generates mycotoxins in wheat grains that can cause serious health problems in humans and livestock (Powell et al., 2017). 相似文献
8.
Rizky Pasthika Kirana Kumar Gaurav Sanu Arora Gerlinde Wiesenberger Maria Doppler Sebastian Michel Simone Zimmerl Magdalena Matic Chinedu E. Eze Mukesh Kumar Ajla Topuz Marc Lemmens Rainer Schuhmacher Gerhard Adam Brande B. H. Wulff Hermann Buerstmayr Barbara Steiner 《Plant biotechnology journal》2023,21(1):109-121
Aegilops tauschii is the diploid progenitor of the wheat D subgenome and a valuable resource for wheat breeding, yet, genetic analysis of resistance against Fusarium head blight (FHB) and the major Fusarium mycotoxin deoxynivalenol (DON) is lacking. We treated a panel of 147 Ae. tauschii accessions with either Fusarium graminearum spores or DON solution and recorded the associated disease spread or toxin-induced bleaching. A k-mer-based association mapping pipeline dissected the genetic basis of resistance and identified candidate genes. After DON infiltration nine accessions revealed severe bleaching symptoms concomitant with lower conversion rates of DON into the non-toxic DON-3-O-glucoside. We identified the gene AET5Gv20385300 on chromosome 5D encoding a uridine diphosphate (UDP)-glucosyltransferase (UGT) as the causal variant and the mutant allele resulting in a truncated protein was only found in the nine susceptible accessions. This UGT is also polymorphic in hexaploid wheat and when expressed in Saccharomyces cerevisiae only the full-length gene conferred resistance against DON. Analysing the D subgenome helped to elucidate the genetic control of FHB resistance and identified a UGT involved in DON detoxification in Ae. tauschii and hexaploid wheat. This resistance mechanism is highly conserved since the UGT is orthologous to the barley UGT HvUGT13248 indicating descent from a common ancestor of wheat and barley. 相似文献
9.
W. D. Bovill M. Horne D. Herde M. Davis G. B. Wildermuth M. W. Sutherland 《TAG. Theoretical and applied genetics. Theoretische und angewandte Genetik》2010,121(1):127-136
Crown rot of wheat (Triticum aestivum), predominantly caused by the fungus Fusarium pseudograminearum, has become an increasingly important disease constraint in many winter cereal production regions in Australia. Our group has previously identified a range of quantitative trait loci (QTL) for partial resistance to crown rot in various bread wheat sources. Here, we report on work that has assessed the effectiveness of pyramiding QTL to improve resistance to crown rot. Two doubled haploid populations were analysed—one from a cross between two previously characterised sources of partial seedling resistance (2-49 and W21MMT70; n = 208) and one from a cross between 2-49 and the commercial variety Sunco, a source of adult field resistance (n = 134). Both populations were phenotyped for seedling resistance to crown rot. Microsatellite and DArT markers were used to construct whole genome linkage maps for use in composite interval mapping (CIM) to identify QTL. Three QTL were detected in both trials conducted on the 2-49/W21MMT70 population. These were located on chromosomes 1D (QCr.usq-1D.1), 3B (QCr.usq-3B.1) and 7A. QCr.usq-1D.1 and the previously undetected 7A QTL were inherited from 2-49. QCr.usq-3B.1, inherited from W21MMT70, was the most significant of the QTL, explaining up to 40.5% of the phenotypic variance. Three QTL were identified in multiple trials of the Sunco/2-49 population. These were located on chromosomes 1D (QCr.usq-1D.1), 2B (QCr.usq-2B.2) and 4B (QCr.usq-4B.1). Only QCr.usq-2B.2 was inherited from Sunco. QCr.usq-4B.1 was the most significant of these QTL, explaining up to 19.1% of the phenotypic variance. In the 2-49/W21MMT70 population, several DH lines performed significantly better than either parent, with the best recording an average disease severity rating of only 3.8% of that scored by the susceptible check cultivar Puseas. These lines represent a new level of seedling crown rot resistance in wheat. 相似文献
10.
H. B. Li W. Yan G. R. Liu S. M. Wen C. J. Liu 《TAG. Theoretical and applied genetics. Theoretische und angewandte Genetik》2011,122(2):395-403
Tan spot, caused by Pyrenophora tritici-repentis, is a foliar disease of wheat, and it can inflict serious reduction in grain yield and quality. The bread wheat variety Ernie was found to be immune to this disease in Australia, and its genetic control was investigated by quantitative trait loci (QTL) analysis using a doubled haploid population. Eight QTL were identified in this population from three independent trials, and four of them were derived from the parent Ernie. The most significant QTL was located on chromosome arm 2BS, explaining 38.2, 29.8 and 36.2% of the phenotypic variance, respectively, in these trials. The effects of the 2BS QTL were further validated in four additional populations. The presence of this single QTL reduced disease severity by between 29.2 and 67.1% with an average of 50.5%. The significant effects of this QTL and its consistent detection across all the trials with different genetic backgrounds make it an ideal target for breeding programmes as well as for its further characterization. Data from this study also showed that neither plant height nor heading date significantly affects tan spot resistance. 相似文献
11.
12.
A major QTL for resistance to Gibberella stalk rot in maize 总被引:1,自引:0,他引:1
Qin Yang Guangming Yin Yanling Guo Dongfeng Zhang Shaojiang Chen Mingliang Xu 《TAG. Theoretical and applied genetics. Theoretische und angewandte Genetik》2010,121(4):673-687
Fusarium graminearum Schwabe, the conidial form of Gibberella zeae, is the causal fungal pathogen responsible for Gibberella stalk rot of maize. Using a BC1F1 backcross mapping population derived from a cross between ‘1145’ (donor parent, completely resistant) and ‘Y331’ (recurrent
parent, highly susceptible), two quantitative trait loci (QTLs), qRfg1 and qRfg2, conferring resistance to Gibberella stalk rot have been detected. The major QTL qRfg1 was further confirmed in the double haploid, F2, BC2F1, and BC3F1 populations. Within a qRfg1 confidence interval, single/low-copy bacterial artificial chromosome sequences, anchored expressed sequence tags, and insertion/deletion
polymorphisms, were exploited to develop 59 markers to saturate the qRfg1 region. A step by step narrowing-down strategy was adopted to pursue fine mapping of the qRfg1 locus. Recombinants within the qRfg1 region, screened from each backcross generation, were backcrossed to ‘Y331’ to produce the next backcross progenies. These
progenies were individually genotyped and evaluated for resistance to Gibberella stalk rot. Significant (or no significant) difference in resistance reactions between homozygous and heterozygous genotypes
in backcross progeny suggested presence (or absence) of qRfg1 in ‘1145’ donor fragments. The phenotypes were compared to sizes of donor fragments among recombinants to delimit the qRfg1 region. Sequential fine mapping of BC4F1 to BC6F1 generations enabled us to progressively refine the qRfg1 locus to a ~500-kb interval flanked by the markers SSR334 and SSR58. Meanwhile, resistance of qRfg1 to Gibberella stalk rot was also investigated in BC3F1 to BC6F1 generations. Once introgressed into the ‘Y331’ genome, the qRfg1 locus could steadily enhance the frequency of resistant plants by 32–43%. Hence, the qRfg1 locus was capable of improving maize resistance to Gibberella stalk rot. 相似文献
13.
14.
Crown rot is one of the main important fungal diseases affecting wheat in many areas of the world, including Australia, USA, and Iran. Until now, there had been no report of this pathogen in Iraq. Plants displaying crown rot symptoms were observed in Shaat Alarab (Basra, Iraq); we investigated the causal agent of the disease. Samples were surface-sterilized in bleach (1% available chlorine) and cultured on quarter-strength potato dextrose agar plates. DNA was extracted from fungal mycelia, using a modified CTAB protocol. The ITS/5.8S regions were amplified using primer pair ITS1 and ITS4. PCR products purified using a gel extraction kit were sequenced. The sequence that was detected was used to BLAST against NCBI data. The most similar sequence was the ITS/5.8S rDNA region of Fusarium pseudograminearum (strain NRRL28062), showing 97.95% identity. This species normally causes crown rot, resulting in severe damage under dry spring conditions. A pathogenicity test employed to assess the disease-causing ability of the strain showed significant disease symptoms up to 57% infected spikelets. The results confirmed the presence of F. pseudograminearum as a causal agent of wheat crown rot in Iraq. The presence of this pathogen demands further investigations to develop resistant cultivars and/or mechanical control. 相似文献
15.
Lowe I Jankuloski L Chao S Chen X See D Dubcovsky J 《TAG. Theoretical and applied genetics. Theoretische und angewandte Genetik》2011,123(1):143-157
A mapping population of 186 recombinant inbred lines developed from a cross between UC1110, an adapted California spring wheat,
and PI610750, a synthetic derivative from CIMMYT’s Wide Cross Program, was evaluated for its response to current California
races of stripe rust (Puccinia striiformis f. sp. tritici) in replicated field trials over four seasons (2007–2010) in the northern Sacramento Valley. A genetic map was constructed
consisting of 1,494 polymorphic probes (SSRs, DArTs, and ESTs) mapped to 558 unique loci, and QTL analysis revealed the presence
of four stripe rust resistance QTL segregating in this population, two from UC1110 (on chromosomes 3BS and 2BS) and two from
PI610750 (5AL and 2AS). The two QTL of largest effects (on 3BS and 5AL) were validated in independent populations and their
intervals narrowed to 2.5 and 5.3 cM, respectively. The 3BS QTL was shown, by allelism test and genotype, to carry a gene
different from the Yr30/Sr2 complex. Mapped position also suggests that the 3BS QTL is associated with a gene different from either Yrns-B1 or YrRub, two stripe rust resistance genes mapped to this region in other studies. The 5AL QTL carries a previously unreported partial
stripe rust resistance gene, designated here as Yr48. This paper discusses the individual contributions to resistance of these four QTL, their epistatic interactions, and their
potential in durable resistance breeding strategies based on combinations of partial resistance genes. 相似文献
16.
Identification and mapping of fruit rot resistance QTL in American cranberry using GBS 总被引:1,自引:0,他引:1
Guillaume Daverdin Jennifer Johnson-Cicalese Juan Zalapa Nicholi Vorsa James Polashock 《Molecular breeding : new strategies in plant improvement》2017,37(3):38
Sustainability of the cranberry industry is threatened by widespread and increasing losses due to fruit rot in the field as well as increasing restrictions on fungicide inputs. Breeding for resistance offers a partial solution but is challenging because fruit rot is caused by a complex of pathogenic fungi that can vary by location and from year to year. We identified four genetically diverse germplasm accessions that exhibit broad-spectrum fruit rot resistance under field conditions. Three of these accessions were used in biparental crosses to develop four populations segregating for resistance. Genotyping by sequencing was used to generate single-nucleotide polymorphism (SNP) markers for development of high-density genetic maps and quantitative trait locus (QTL) analyses. Nineteen QTL associated with fruit rot resistance, distributed on nine linkage groups, were discovered in our populations. Three of these QTL matched previously reported fruit rot resistance QTL. Four newly reported QTL found on linkage group 8 (Vm8), which explain between 21 and 33% of the phenotypic variance for fruit rot, are of particular interest to our breeding program. The populations described herein were also phenotyped for other horticulturally important traits, and QTL associated with yield and berry weight were identified. These QTL provide markers for candidate gene discovery and for future breeding efforts to enhance and pyramid disease resistance and other traits into elite horticultural backgrounds. 相似文献
17.
Guo PG Bai GH Shaner GE 《TAG. Theoretical and applied genetics. Theoretische und angewandte Genetik》2003,106(6):1011-1017
Large-scale field screening for Fusarium head blight (FHB) resistance in wheat is difficult because environmental factors strongly influences the expression of resistance genes. Marker-assisted selection (MAS) may provide a powerful alternative. Conversion of amplified fragment length polymorphism (AFLP) markers into sequence-tagged site (STS) markers can generate breeder-friendly markers for MAS. In a previous study, one major quantitative trait locus (QTL) on chromosome 3BS was identified by using EcoRI-AFLP and a recombinant inbred population derived from the cross Ning 7840/Clark. Further mapping with PstI-AFLPs identified five markers that were significantly associated with the QTL. Three of them individually explained 38% to 50% of the phenotypic variation for FHB resistance. Two of them (pAGT/mCTG57, pACT/mCTG136) were linked to the QTL in coupling, and another (pAG/mCAA244) was linked to the QTL in repulsion. Successful conversion of one AFLP marker (pAG/mCAA244) yielded a co-dominant STS marker that explains about 50% of the phenotypic variation for FHB resistance in the population. The STS was validated in 14 other cultivars and is the first STS marker for a FHB resistance QTL converted from an AFLP marker. 相似文献
18.
Katherine J. Linsell Muhammad S. Rahman Julian D. Taylor Rowena S. Davey Beverley J. Gogel Hugh Wallwork Kerrie L. Forrest Matthew J. Hayden Sharyn P. Taylor Klaus H. Oldach 《TAG. Theoretical and applied genetics. Theoretische und angewandte Genetik》2014,127(6):1409-1421
Key message
A whole genome average interval mapping approach identified eight QTL associated with P. thornei resistance in a DH population from a cross between the synthetic-derived wheat Sokoll and cultivar Krichauff.Abstract
Pratylenchus thornei are migratory nematodes that feed and reproduce within the wheat root cortex, causing cell death (lesions) resulting in severe yield reductions globally. Genotypic selection using molecular markers closely linked to Pratylenchus resistance genes will accelerate the development of new resistant cultivars by reducing the need for laborious and expensive resistance phenotyping. A doubled haploid wheat population (150 lines) from a cross between the synthetic-derived cultivar Sokoll (P. thornei resistant) and cultivar Krichauff (P. thornei moderately susceptible) was used to identify quantitative trait loci (QTL) associated with P. thornei resistance. The resistance identified in the glasshouse was validated in a field trial. A genetic map was constructed using Diversity Array Technology and the QTL regions identified were further targeted with simple sequence repeat (SSR) and single-nucleotide polymorphism (SNP) markers. Six significant and two suggestive P. thornei resistance QTL were detected using a whole genome average interval mapping approach. Three QTL were identified on chromosome 2B, two on chromosome 6D, and a single QTL on each of chromosomes 2A, 2D and 5D. The QTL on chromosomes 2BS and 6DS mapped to locations previously identified to be associated with Pratylenchus resistance. Together, the QTL on 2B (QRlnt.sk-2B.1–2B.3) and 6D (QRlnt.sk-6D.1 and 6D.2) explained 30 and 48 % of the genotypic variation, respectively. Flanking PCR-based markers based on SSRs and SNPs were developed for the major QTL on 2B and 6D and provide a cost-effective high-throughput tool for marker-assisted breeding of wheat with improved P. thornei resistance. 相似文献19.
Saturation and comparative mapping of a major Fusarium head blight resistance QTL in tetraploid wheat 总被引:2,自引:0,他引:2
Xunfen Chen Justin D. Faris Jinguo Hu Robert W. Stack Tika Adhikari Elias M. Elias Shahryar F. Kianian Xiwen Cai 《Molecular breeding : new strategies in plant improvement》2007,19(2):113-124
Fusarium head blight (FHB) is a devastating disease of cultivated wheat worldwide. Partial resistance to FHB has been identified
in common wheat (Triticum aestivum L.). However, sources of effective FHB resistance have not been found in durum wheat (T. turgidum L. var. durum). A major FHB resistance quantitative trait loci (QTL), Qfhs.ndsu-3AS, was identified on chromosome 3A of T.
dicoccoides, a wild relative of durum wheat. Here, we saturated the genomic region containing the QTL using EST-derived target region
amplified polymorphism (TRAP), sequence tagged site (STS), and simple sequence repeat (SSR) markers. A total of 45 new molecular
marker loci were detected on chromosome 3A and the resulting linkage map consisted of 55 markers spanning a genetic distance
of 277.2 cM. Qfhs.ndsu-3AS was positioned within a chromosomal interval of 11.5 cM and is flanked by the TRAP marker loci, Xfcp401 and Xfcp397.2. The average map distance between the marker loci within this QTL region was reduced from 4.9 cM in the previous study to
3.5 cM in the present study. Comparative mapping indicated that Qfhs.ndsu-3AS is not homoeologous to Qfhs.ndsu-3BS, a major FHB QTL derived from the common wheat cultivar Sumai 3. These results facilitate our efforts toward map-based cloning
of Qfhs.ndsu-3AS and utilization of this QTL in durum wheat breeding via marker-assisted selection. 相似文献
20.
Genqiao Li Ying Wang Ming-Shun Chen Erena Edae Jesse Poland Edward Akhunov Shiaoman Chao Guihua Bai Brett F Carver Liuling Yan 《BMC genomics》2015,16(1)