摘要
小麦种子储藏蛋白主要包括谷蛋白和醇溶蛋白,它们决定小麦独特的加工品质。醇溶蛋白有四种类型,即α-、γ-、ω-和LMW-醇溶蛋白,占小麦种子储藏蛋白总量的40-50%,是人类日常摄取所需蛋白营养的主要来源之一。因此醇溶蛋白影响面粉的加工品质和营养品质。小麦近缘属种中含有丰富的醇溶蛋白基因资源。赤霉病是世界小麦三大主要病害之一,其主要致病菌是禾谷镰刀菌(Fusarium graminearum)。小麦感染赤霉病后不仅产量降低、种子储藏蛋白的合成受到抑制,而且会被赤霉菌分泌的真菌毒素污染,对食品加工和饲料行业造成严重威胁。因此世界各主要小麦生产国对赤霉菌毒素含量都有限制性标准。本文对醇溶蛋白进行了基因克隆、结构分析、二硫键形成方式和进化等方面的研究,并探索了植物激素与小麦赤霉病抗性之间的关系。主要结果如下
1.从野生二粒小麦中克隆了4个α-醇溶蛋白基因(DQ140349-DQ140351)的编码序列。这些序列全长约800 bp,没有内含子。推测DQ140351和DQ140352在胚乳中可以正常表达,而DQ140349和DQ140350是假基因。忽略提前终止密码子和移码突变,得到这4条基因的推导氨基酸序列：它们均含有6个保守的半胱氨酸：重复区的重复单元与已报导的结果有区别；乳糜泻抗原主要分布于成熟蛋白N末端；脯氨酸密码子的使用有明显偏好；谷氨酰胺在整个蛋白中含量丰富,但在特定区域呈现集中分布的特点。
2.从小麦中克隆了10条新型α-醇溶蛋白基因(Gli-ta、Gli-turgl-6和Gli-cs1-3)其中Gli-csl、Gli-cs2、Gli-cs3和Gli-turg6是假基因。这些新型α-醇溶蛋白具有显著不同于典型α-醇溶蛋白的特征。推测这些新型α-醇溶蛋白基因起源于发生在典型α-醇溶蛋白基因两个多聚谷氨酰胺区之间的非常规重组。与典型α-醇溶蛋白相比,Gli-cs3的分子量更大且有10个半胱氨酸残基,它的一级结构多一个特征区I和一个多聚谷氨酰胺区；其余9条序列缺失一个特征区1和一个多聚谷氨酰胺区,其推导蛋白的分子量较小且仅有2个半胱氨酸残基。Gli-talGli-ta-like a-醇溶蛋白基因在种子发育过程中能正常表达。SDS-PAGE分析表明Gli-ta的2个半胱氨酸都能参与形成分子间二硫键；回收种子提取物中与Gli-ta具有相似迁移率的条带并进行质谱分析,结果表明该条带中包含Gli-ta/Gli-ta-like a-醇溶蛋白,而且它们在种子内参与形成分子间二硫键。
3.从Genbank中下载了79条节节麦α-醇溶蛋白基因序列和40条已知染色体来源的α-醇溶蛋白基因序列用于开发6D染色体特异的α-醇溶蛋白基因的SNP标记。总共设计了85条SNP标记引物用于检测68个潜在的6D染色体特异的α-醇溶蛋白基因的SNP位点。以中国春第6部分同源群染色体的缺体-四体材料和六倍体小麦的二倍体和四倍体祖先种的基因组DNA为PCR模板,验证得到11个6D染色体特异的SNP标记。以1份人工合成六倍体小麦及其亲本DNA为PCR模板,证明在节节麦和圆锥小麦杂交形成六倍体小麦的过程中六倍体小麦基因组会丢失部分α-醇溶蛋白基因。以节节麦、普通小麦及其4个亚种的DNA为PCR模板,证明至少两种遗传背景不同的节节麦参与了普通小麦D基因组的形成。
4.从普通小麦及其近缘物种中克隆了170条γ-醇溶蛋白基因序列,其中138条是潜在的功能基因。这些基因开放阅读框的长度介于678-1089 bp。基因之间长度的差异主要是因为它们重复区的长度不一致,即重复单元(P(Q/L/S/T/I/V/WA)F(S/Y/V/Q/I/C/L)P (R/L/S/T/H/C/Y)Q1-2(P(S/L/T/A/F/H)QQ)1-2)的数目多少不一。序列多样性和连锁不平衡分析表明Y-醇溶蛋白基因家族具有非常高的遗传多样性。系统进化分析表明节节麦和斯卑尔脱山羊草组(Siptopsis)物种的γ-醇溶蛋白基因差异不显著(即相比于A基因组,B(S)和D基因组的γ-醇溶蛋白基因的亲缘关系更近)。根据半胱氨酸数目和相对位置的不同,可将Y-醇溶蛋白基因分为9种类型(C1-9)和17(SG-1至SG-17)个亚类；根据重复区长度的差异,γ-醇溶蛋白可被分为两类。氨基酸组成分析表明人体所需的8种必需氨基酸含量在γ-醇溶蛋白中差异巨大。SG-10.SG-12和重复区较短的γ-醇溶蛋白必需氨基酸含量较高。C9类和重复区较短的γ-醇溶蛋白几乎不含有乳糜泻抗原表位。不同γ-醇溶蛋白类群的乳糜泻抗原含量和必需氨基酸含量不同。因此下一步研究应当更重视重复区较短的γ-醇溶蛋白,其营养品质更高且几乎不含乳糜泻抗原。这为培育品质优良、无乳糜泻毒性的小麦品种提供了可能。
5.从小麦和山羊草属物种中克隆了4条新型Y-醇溶蛋白基因(Gli-ngl-Gli-ng4),其中Gli-ng1、Gli-ng2和Gli-ng3的核酸序列完全一致。与典型Y-醇溶蛋白相比,新型Y-醇溶蛋白的分子量相对较小,其一级结构缺失非重复区、谷氨酰胺富集区、重复区的3’端和C-末端区的5’端。依据核酸序列特征,我们推测这些新型γ-醇溶蛋白基因起源于发生在典型Y-醇溶蛋白基因重复区和C-末端区之间的非常规重组,并找到了其可能的祖先基因。典型γ-醇溶蛋白具有8个或9个半胱氨酸,而Gli-ng1和Gli-ng4仅分别含有2个和3个半胱氨酸。cDNA克隆表明,Gli-ng1能在中国春种子发育过程中正常转录。SDS-PAGE分析表明Gli-ng1的2个半胱氨酸在体外均可参与形成分子间二硫键。系统进化分析表明这4条新型γ-醇溶蛋白基因与来源于小麦B基因组或山羊草S基因组的Y-醇溶蛋白基因亲缘关系最近。
6.首次克隆得到了大麦和黑麦α-醇溶蛋白的基因序列。结构特征分析表明它们与Gli-ta/Gli-ta-like新型α-醇溶蛋白基因具有相似的结构,这说明非常规重组可能导致α-醇溶蛋白基因从黑麦和大麦基因组中丢失了。首次得到黑麦40Kγ-secalin基因序列,结构分析表明它们与典型γ-醇溶蛋白基因具有高度同源性,而且由于75Kγ-secalin的重复区比40Kγ-secalin长,因此分子量较大。系统分析了小麦族二倍体物种707个α-醇溶蛋白基因和280个γ-醇溶蛋白基因的多样性。根据一级结构特点,可将α-和γ-醇溶蛋白分别分为4和4类。不同类群醇溶蛋白的一级结构差异很大。根据半胱氨酸数目和相对位置的差异,可将具有潜在功能和完整开放阅读框的225条α-醇溶蛋白序列和168条γ-醇溶蛋白基因分别分为15和17种类型。一些α-和γ-醇溶蛋白包含奇数个半胱氨酸,这表明它们有可能形成分子间二硫键,因此有可能在种子内参与谷蛋白聚合物的形成。
7.水杨酸在固体和液体培养基中能显著抑制F. graminearum菌丝的生长及孢子的萌发,并随着水杨酸浓度的提高最终杀死病菌；水杨酸在抑制菌丝体生长的同时也抑制F. graminearum合成真菌毒素DON；酸性条件下水杨酸对F. graminearum的抑制作用较强,在碱性条件下F. graminearum能使用水杨酸作为唯一碳源。高效液相色谱分析进一步证明F. graminearum具有代谢水杨酸的能力。为深入了解水杨酸对F. graminearum菌丝生长的抑制作用,我们比较了在液体培养基中添加水杨酸8h和24 h后F. graminearum基因在菌丝中的表达情况,结果表明F. graminearum能通过儿茶酚和/或龙胆酸途径代谢水杨酸。因此,虽然高浓度的水杨酸可直接影响孢子萌发率和菌丝生长速率,但F.graminearum具有代谢水杨酸的能力。将水杨酸与F. graminearum孢子混合后接种极易感染赤霉菌的小麦品种'Roblin'能显著减轻其赤霉病的症状；但若先接种水杨酸后接种孢子或者先接种孢子再接水杨酸,赤霉病的症状与对照没有显著差异。通过检测PR1、NPR1、Pdf1.2和PR4等与植物防御反应相关基因的表达情况,证明水杨酸诱导的抗病反应对提高小麦赤霉病抗性无显著作用。
8.小麦穗子被F. graminearum侵染会累积水杨酸(SA)、茉莉酸(JA)、脱落酸(ABA)及其代谢物和生长素(IAA)及其缀合物。F. graminearum能合成IAA、少许ABA及其代谢物是这些激素在穗子中含量增加的原因之一。小麦穗子的激素反应网络显示SA刺激JA的累积,但不影响ABA和IAA；JA能增加ABA含量,降低SA水平,但对IAA无影响：施加ABA导致JA和IAA水平的提高及SA水平的降低；施加IAA引起ABA浓度的增加,但不影响SA和JA；12种受检赤霉素的含量均低于检测水平。JA和IAA可显著抑制F. graminearum菌丝的生长和孢子的萌发,而ABA只抑制菌丝的生长。在改良SNA平板上,IAA在抑制菌丝蔓延的同时能显著增加菌丝体的厚度。将JA或IAA与孢子混合后接种能显著减轻赤霉病症状。若不将JA/IAA与孢子在接种前混合,它们对赤霉病的抑制效果明显减弱。接种ABA显著降低小麦对赤霉病的抵抗力。除去植物激素对F. graminearum的直接影响,我们发现SA不影响,甚至对赤霉病有轻微的促进作用；JA和IAA显著抑制赤霉病发病水平；ABA促进赤霉病的发生和发展,但不影响F. graminearum的生长,同时意外的发现ABA显著抑制染病穗子积累真菌毒素DON。降低小麦穗子JA和ABA的水平证明它们分别能增强和减弱小麦对F. graminearum的抗性水平。通过检测ABT2、(3-expansin、LEA、PR1、NPR1、Pdfl.2和PR4等与植物激素信号途径相关基因的表达,证明SA与JA相互拮抗控制PRl基因的表达,而JA与ABA在拮抗调控小麦赤霉病抗性的同时,相互协作控制一些抗性相关基因的表达。
The unique properties of wheat flour primarily depend on seed storage proteins, one of the most important sources of protein for human being, which mainly consist of glutenins and gliadins. Gliadins can be classified as four groups, i.e.α-,γ-,ω-and LMW-gliadins, and account for about 40-50% of the total seed storage proteins. Gliadins significantly affect the processing and nutritional qualities of flour. There is a great diversity of gliadin genes in wheat and its closely related species. Fusarium head blight (FHB) is one of the most common diseases on wheat worldwide. FHB can cause significant yield losses and quality reductions. Fusarium graminearum is the principal causal agent of FHB, and produces the mycotoxin deoxynivalenol (DON). DON is a potent inhibitor of protein synthesis, and thereby presents hazards for both humans and animal health. Therefore, many countries have adopted maximum allowable limits for the mycotoxin DON in grain. This research focused on gene cloning, structural analysis, disulphide bond formation and gene evolution of gliadins, and investigated the role of salicylic acid on wheat FHB resistance. The main results are as follows:
1. The coding regions of four a-gliadin genes of wild emmer wheat (GenBank No. DQ140349-DQ140352) were obtained. Each was about 800 bp long, and intronless. DQ140351 and DQ140352 were potentially functional in endosperm, whereas DQ140349 and DQ140350 were both pseudogenes. Alignment of their deduced amino acid sequences (by removing the in-frame stop condons and frameshift mutation) indicated that six cysteine residues were not randomly distributed, but conserved in the two unique domains; The motif unit of the repetitive domain for the four newly-detected genes was different from previous reports; the N-terminal part of a-gliadins was more toxic to celiac patients; codon usage for proline was biased; codons for glutamine were clustered into specific regions.
2. Ten novel a-gliadin genes (Gli-ta, Gli-turgl-6 and Gli-csl-3) with unique characteristics were isolated from wheat, among which Gli-cs1, Gli-cs2, Gli-cs3 and Gli-turg6 were pseudogenes. Gli-cs3 and nine other sequences were much larger and smaller, respectively, than the typical a-gliadins. This variation was caused by insertion or deletion of the unique domain I and a polyglutamine region, possibly due to illegitimate recombination. Consequently, Gli-cs3 contained ten cysteine residues, while there were only two cysteine residues in the other nine sequences. Gli-ta/Gli-ta-like a-gliadin genes are normally expressed during the development of seeds. SDS-PAGE analysis showed that in vitro expressed Gli-ta could form intermolecular disulphide bonds, and act as a chain extender. A protein band similar in size to Gli-ta has been observed in seed extracts, and mass spectrometry results confirm that the band contains small molecular weight a-gliadins, which is a characteristic of the novel a-gliadins. Mass spectrometry results also indicated that the two cysteine residues of Gli-ta/Gli-ta-like proteins participated in the formation of intermolecular disulphide bonds in vivo.
3. Seventy-nine a-gliadin sequences cloned from Aegilops lauschii and another 40 a-gliadin genes with known chromosome locations were downloaded from Genbank and used to design chromosome 6D-specific SNP markers for a-gliadin genes. A total of 85 SNP primers were designed to detect 68 candidate chromosome 6D-specific SNPs. Experimental tests revealed 11 chromosome 6D-specific SNP markers by using genomic DNA from homoeologous group-6 nullisomic-tetrasomic lines of Chinese Spring and putative diploid and tetraploid ancestors of hexaploid wheat as PCR templates. Detection of SNP markers in one synthetic hexaploid wheat and its parental lines indicated that some a-gliadins genes were lost from the Gli-2 loci during the formation of hexaploid wheat by amphidiploidization of the genomes of Triricum turgidum and Aegilops tauschii. Detection of these SNP markers in Ae. tauschii, Triticum aestivum and its four subspecies indicated that at least two genetically distinct sources of Ae. tauschii contributed germplasm to the D genome of T. aestivum.
4. A total of 170 y-gliadin genes were isolated from common wheat and its closely related species, among which 138 sequences were putatively functional. The ORF length of these sequences ranged from 678 to 1089 bp, and the repetitive region is mainly responsible for the size heterogeneity ofγ-gliadins. The motif P(Q/L/S/T/I/V/R/A)F(S/Y/V/Q/I/C/L)P(R/L/S/ T/H/C/Y)Q,_2(P(S/L/T/A/F/H)QQ)1-2 is repeated from 7 to 22 times. Sequence polymorphism and linkage disequilibrium analyses show that y-gliadins are highly diverse. Phylogenic analyses indicated that there is no obvious discrimination between Sitopsis and Ae. tauschii at the Gli-1 loci, compared with diploid wheat. According to the number and placement of cysteine residues, we defined nine cysteine patterns and 17 subgroups. Alternatively, we classifiedγ-gliadins into two types based on the length of the repetitive domain. Amino acid composition analyses indicate that there is a wide range of essential amino acids inγ-gliadins, and thoseγ-gliadins from subgroup SG-10 and SG-12 and y-gliadins with a short repetitive domain are more nutritional. A screening of toxic epitopes shows that y-gliadins with a pattern of C9 and y-gliadins with a short repetitive domain almost lack any epitopes. It is suggested that the genes with a short repetitive domain are more nutritional and valuable. Therefore, each group/subgroup contributes differently to nutritional quality and epitope content, and it would be possible to breed wheat varieties containing the y-gliadins that are less, even non-toxic and more nutritional.
5. Four novel y-gliadin genes (Gli-ngl to Gli-ng4) were cloned from wheat(Triticum aestivum) and Aegilops species. The novelγ-gliadins were much smaller in molecular size when compared to the typical y-gliadins, which was caused by deletion of the non-Repetitive domain, Glutamine-rich region,3'part of the Repetitive domain and 5'part of the C-terminal, possibly due to illegitimate recombination between the repetitive domain and the C-terminal. As a result, Gli-ngl and Gli-ng4 only contained two and three cysteine residues, respectively. Gli-ngl, as the representative of novel y-gliadin genes, has been sub-cloned into an E. coli expression system. SDS-PAGE indicated that both cysteine residues of Gli-ngl could participate in the formation of intermolecular disulphide bonds in vitro. Successful cloning of Gli-ng1 from seed cDNA of T. aestivum cv'Chinese Spring' suggested that these novel y-gliadin genes were normally transcribed during the development of seeds. Phylogenic analysis indicated that the four novelγ-gliadin genes had a closer relationship with those from the B(S) genome of wheat.
6. Alpha-gliadin genes with unique structure originating from illegitimate recombination were obtained from barley and rye, indicating that the absence of a-gliadin genes in barley and rye is a result of illegitimate recombination. Analysis of the full coding region of 40K y-secalin genes indicated that they have a very close structural relationship to the typical y-gliadin genes of wheat, and that the higher molecular weight of 75K y-secalins than that of 40K y-secalins is a result of the addition of repetitive sequence. Molecular diversity ofα-and y-prolamin genes was systematically investigated. According to their distinct constitutions of protein domains, we classifyα-andγ-prolamins into 4 and 4 groups. According to the number and placement of cysteine residues, we defined 15 and 17 cysteine patterns for a-and y-prolamins, respectively. Some of theα-andγ-prolamins contain odd numbers of cysteine residues, which strongly suggests that they possibly participate in the formation of intermolecular disulphide bonds.
7. The mycelial growth and spore germination of F. graminearum were significantly inhibited, and eventually halted by the presence of increasing concentration of SA in both liquid and solid media. Addition of SA also significantly reduced the production of the mycotoxin deoxynivalenol (DON). However the inhibitory effect of SA required acidic growth conditions to be observed while basic conditions allowed F. graminearum to use SA as a sole carbon source. HPLC analysis confirmed the capacity of F. graminearum to metabolize SA. To better understand the effect of SA on F. graminearum mycelial growth, we have compared the expression profiles of SA-treated and untreated F. graminearum liquid cultures after 8 and 24 h of treatment, using an F. graminearum custom microarray. The microarray analysis suggested that F. graminearum can metabolize SA through either the catechol or gentisate pathways that are present in some fungal species. Inoculation of F. graminearum spores in a SA-containing solution has led to reduced FHB symptoms in the very susceptible Triticum aestivum cultivar Roblin. In contrast, no inhibition was observed when SA and spores were inoculated sequentially. The expression patterns for the wheat PR1, NPR1, Pdfl.2 and PR4 genes, a group of indicator genes for the defence response, suggested that SA-induced resistance contributed little to the reduction of symptoms in our assay conditions. Our results demonstrates that, although F. graminearum has the capacity to metabolize SA, SA has a significant and direct impact on F. graminearum through a reduction in efficiency of germination and growth at higher concentrations.
8. F. graminearum challenge induced the accumulation of SA, JA, ABA and its metabolites, IAA and IAA conjugates in wheat heads, partially because F. graminearum is able to produce IAA, and a little bit ABA and its metabolites. Testing of the hormone networking showed that application of SA promoted the accumulation of JA, whereas it did not affect ABA and IAA levels; application of JA increased the amount of ABA and reduced the level of SA, whereas did not affect IAA; application of ABA resulted in an increase in the JA and IAA levels and a decrease in the SA level; application of IAA induced an increased concentration of ABA, whereas it had no affect on SA and JA; GAs monitored were not quantifiable in those assays. The growth of F. graminearum mycelia and the germination of spores could be significantly inhibited by JA and IAA, whereas ABA only suppressed mycelial growth within the tested concentrations. IAA significantly increased density of mycelia on modified SNA plates, while inhibiting the spread of mycelia. Inoculation of F. graminearum spore suspensions containing either JA or IAA-containing solution led to reduced visual FHB symptoms. In contrast, less inhibition was observed when JA or IAA and spores were inoculated sequentially. Inoculation of ABA in heads enhanced susceptibility to FHB. To further understand the effect of hormones on FHB resistance, we sequentially inoculated hormones and spores into each spikelet of a spike. As a result, SA did not affect, and even slightly increased FHB severity; JA and IAA significantly reduced FHB levels; ABA promoted the visual disease symptoms, while unexpectedly suppressing the accumulation of mycotoxin without affecting fungal biomass. Down-regulation of endogenous JA and ABA levels confirmed that JA and ABA respectively functioned as a positive and a negative regulator of wheat defense response to F. graminearum infection. The expression patterns for the wheat ABT2,β-expansin, LEA, PR], NPR1, Pdfl.2 and PR4 genes, a group of indicator genes for the defence response and hormone signaling, suggested that SA antagonistically interacted with JA to control the expression of PR1, and synergistic interactions existed between JA and ABA, even though they antagonistically modulated FHB resistance in wheat.
引文
1. Abrams SR, Nelson K, Ambrose SJ. Deuterated abscisic acid analogs for mass spectrometry and metabolism studies. Journal of Labelled Cmpounds and Radiopharmaceuticals,2003,46:273-283
2. Adie B, Chico JM, Rubio-Somoza 1, et al. Modulation of plant defenses by ethylene. Journal of Plant Growth Regulation,2007,26:160-177
3. Agrawal GK, Tamogami S, Han O, et al. Rice octadecanoid pathway. Biochemical and Biophysical Research Communications,2004,317:1-15
4. Ahmadian A, Gharizadeh B, O'Meara D, et al. Genotyping by apyrase-mediated allele-specific extension. Nucleic Acids Research,2001,29:e121
5. Akhunov ED, Goodyear AW, Geng S, et al. The organization and rate of evolution of wheat genomes are correlated with recombination rates along chromosome arms. Genome Research,2003,13:753-763
6. Aleandri MP, Magro P, Chilosi G. Modulation of host pH during the wheat Fusarium culmorum interaction and its influence on the production and activity of pectolytic enzymes. Plant Pathology,2007,56:517-525
7. Ali B, Sabri AN, Ljung K, et al. Auxin production by plant associated bacteria:impact on endogenous IAA content and growth of Triticum aestivum L. Letters in Applied Microbiology,2009,48:542-547
8. Ali-Benali MA, Alary R, Joudrier P, et al. Comparative expression of five Lea genes during wheat seed development and in response to abiotic stresses by real-time quantitative RT-PCR. Biochimica et Biophysica Acta,2005,1730:56-65
9. Anderson OD, Litts JC, Gautier MF, et al. Nucleic acid sequence and chromosome assignment of a wheat storage protein gene. Nucleic Acids Research,1984, 12:8129-8144
10. Anderson OD, Litts JC, Greene FC. The a-gliadin gene family. I. characterization of ten new wheat a-gliadin genomic clones, evidence for limited sequence conservaton of flanking DNA, and southern analysis of the gene family. Theoretical and Applied Genetics,1997,95:50-58
11. Anderson OD, Greene FC. The a-gliadin gene family. Ⅱ. DNA and protein sequence variation, subfamily structure, and origins of pseudogenes. Theoretical and Applied Genetics,1997,95:59-65
12. Anderson OD, Hisa CC, Adalsteins AE, et al. Identification of several new classes of low-molecular-weight wheat gliadin-related proteins and genes. Theoretical and Applied Genetics,2001a,103:307-315
13. Anderson OD, Hisa CC, Torres V. The wheat γ-gliadin genes:characterization of ten new sequences and further understanding of γ-gliadin gene family structure. Theoretical and Applied Genetics,2001b,103:323-330
14. Anderson JA, Stack RW, Liu S, et al. DNA markers for Fusarium head blight resistance QTLs in two wheat populations. Theoretical and Applied Genetics,2001c, 102:1164-1168
15. Anderson JP, Badruzsaufari E, Schenk PM, et al. Antagonistic interaction between abscisic acid and jasmonic-ethylene signalling pathways modulates defense gene expression and disease resistance in Arabidopsis. Plant Cell,2004,16:3460-3479
16. Arentz-Hansen EH, McAdam SN, Molberg O, et al. Production of a panel of recombinant gliadins for the characterisation of T cell reactivity in coeliac disease. Gut 2000,46:46-51
17. Arentz-Hansen H, McAdam SN, Molberg O, et al. Celiac lesion T cells recognize epitopes that cluster in regions of gliadins rich in proline residues. Gastroenterology, 2002,123:803-809
18. Bai GH. Deocynivalenol-nonproducing Fusarium graminearum causes initial infection, but does not cause disease spread in wheat spikes. Mycopathologia,2001, 153:91-98
19. Bai G, Shaner G. Management and resistance in wheat and barley to Fusarium head blight. Annual Review of Phytopathology,2004,42:135-161
20. Bari R, Jones JDG. Role of plant hormones in plant defence responses. Plant Molecular Biology,2009,69:473-488
21. Bartels D, Altosaar I, Harberd NP, et al. Molecular analysis of gamma-gliadin gene families at the complex Gli-1 locus of bread wheat(T. aestivum L.). Theoretical and Applied Genetics,1986,72:845-853
22. Bertini L, Leonardi L, Caporale C, et al. Pathogen-responsive wheat PR4 genes are induced by activators of systemic acquired resistance and wounding. Plant Science, 2003,164:1067-1078
23. Bietz JA, Huebner FR, Sanderson JE, Wall JS. Wheat gliadin homology revealed through N-terminal amino acid sequence analysis. Cereal Chemistry,1977, 54:1070-1083
24. Bietz JA. Cereal prolamin evolution and homology revealed by sequence analysis. Biochemical Genetics,1982,20:1039-1053
25. Blanco A, Resta P, Simeone R, et al. Chromosomal location of seed storage protein genes in the genome of Dasypyrum villosum (L.) Candargy. Theoretical and Applied Genetics,1991,82:358-362
26. Buerstmayr H, Lemmens M, Hartl L, et al. Molecular mapping of QTLs for Fusarium head blight resistance in spring wheat.I. Resistance to fungal spread (type Ⅱ resistance). Theoretical and Applied Genetics,2002,104:84-91
27. Bulleid NJ, Freedman RB. The transcription and translation in vitro of individual cereal storage-protein genes from wheat (Triticum aestivum cv. Chinese Spring). Biochemical Journal,1988,254:805-810
28. Bushuk W, Zillman RR. Wheat cultivar identification by gliadin electrophoregrams. I. Apparatus, method, and nomenclature. Canadian Journal of Plant Science,1978, 58:505-515
29. Cain RB, Bilton RF, Darrah JA. The metabolism of aromatic compounds by microorganisms:metabolic pathways in fungi. Biochemical Journal,1968, 108:797-827
30. Caldwell KS, Dvorak J, Lagudah ES, et al. Sequence polymorphism in polyploid wheat and their D-genome diploid ancestor. Genetics,2004,167:941-947
31. Champe SP, El-zayat AAE. Isolation of a sexual sporulation hormone from Aspergillus nidulans. Journal of Bacteriology,1989,171:3982-3988
32. Chen XW, Steed A, Harden C, Nicholson P. Characterization of Arabidopsis thaliana-Fusarium graminearum interactions and identification of variation in resistance among ecotypes. Molecular Plant Pathology,2006,7:391-403
33. Chen X, Steed A, Travella S, et al. Fusarium graminearum exploits ethylene signalling to colonize dicotyledonous and monocotyledonous plants. New Phytologist. 2009,182:975-983
34. Chilosi G, Magro P. Pectolytic enzymes produced in vitro and during colonization of melon tissues by Didymella bryoniae. Plant Patholology,1998,47:700-705
35. Chiwocha SDS, Abrams SR, Ambrose SJ, et al. A method for profiling classes of plant hormones and their metabolites using liquid chromatography-electrospray ionization tandem mass spectrometry:analysis of hormone regulation of thermodormancy of lettuce (Lactuca saliva L.) seeds. Plant Journal,2003,35:405-417
36. Chiwocha SDS, Cutler AJ, Abrams SR, et al. The etrl-2 mutation in Arabidopsis thaliana affects the abscisic acid, auxin, cytokinin and gibberellin metabolic pathways during maintenance of seed dormancy, moist-chilling and germination. Plant Journal, 2005,42:35-48
37. Chono M, Honda I, Schinoda S, et al. Field studies on the regulation of abscisic acid content and germinability during grain development of barley:molecular and chemical analysis of pre-harvest sprouting. Journal of Experimental Botany,2006, 57:2421-2434
38. Ciaffi M, Lee YK, Tamas L, et al. The low-molecular-weight glutenin subunit proteins of primitive wheats. III. The genes from D-genome species. Theoretical and Applied Genetics,1999,98:35-148
39. Clarke BC, Phongkham T, Gianibelli MC, et al. The characterization and mapping of a family of LMW-gliadin genes:effects on dough properties and bread volume. Theoretical and Applied Genetics,2003,106:629-635
40. Cowger C, Patton-Ozkurt J, Brown-Guedira G, et al. Post-anthesis moisture increased Fusarium head blight and deoxynivalenol levels in North Carolina winter wheat. Phytophathology,2009,99:320-327
41. Craig D, Robins G, Howdle PD. Advances in celiac disease. Current Opinion in Gastroenterology,2007,23:142-148
42. Dathe W, Kramell HM, Daeter W, et al. Uptake of jasmonic acid and related compounds by mesophyll protoplasts of the barley leaf. Journal of Plant Growth Regulation,1993,12:133-140
43. De Vos M, Van Oosten VR, Van Poecke RMP, et al. Signal signature and transcriptome changes of Arabidopsis during pathogen and insect attack. Molecular Plant-Microbe Interactions,2005,18:923-937
44. Delaney TP, Uknes S, Vernooij B, et al. A central role of salicylic acid in plant disease resistance. Science,1994,266:1247-1250
45. Desjardins AE, Hohn TM, McCormick SP. Trichothecene biosynthesis in Fusarium: Chemistry, genetics, and significance. Microbiological Reviews 1993,57:595-604
46. Desjardins AE, Proctor RH, Bai G, et al. Reduced virulence of trichothecene-nonproducing mutants of Gibberella zeae in wheat field tests. Molecular Plant-Microbe Interactions,1996,9:775-781
47. Dieterich W, Esslinger B, Schuppan D. Pathomechanisms in celiac disease. International Archives of Allergy and Immunology,2003,132:98-108
48. Ding L. Xu H, Yi H, et al. Resistance to hemi-biotrophic F. graminearum infection is associated with coordinated and ordered expression of diverse defense signalling pathways. PLos one,2011,6:e 19008
49. Dodge AG, Wackett LP. Metabolism of bismuth subsalicylate and intracellular accumulation of bismuth by Fusarium sp. strain BI. Applied and Environmental Microbiology,2005,71:876-882
50. D'Ovidio R, Simeone M, Masci S, et al. Nucleotide sequence of a gamma-type glutenin gene from a durum wheat:Correlation with a gamma-type glutenin subunit from the same biotype. Cereal Chemistry,1995,72:443-449
51. Doyle JJ, Doyle JL. A rapid DNA isolation procedure from small quantities of fresh leaf tissues. Phytochemical Bulletin,1987,19:11-15
52. DuPont FM, Vensel W, Encarnacao T, et al. Similarities of omega gliadins from Triticum urartu to those encoded on chromosome 1A of hexaploid wheat and evidence for their post-translational processing. Theoretical and Applied Genet tics,2004, 108:1299-1308
53. Dvorak J, Luo MC, Yang ZL, Zhang HB. The structure of the Aegilops lauschii genepool and the evolution of hexaploid wheat. Theoretical and Applied Genetics, 1998,97:657-670
54. Elmorjani K, Thievin M, Michon T, et al. Synthetic genes specifying periodic polymers modeled on the repetitive domain of wheat gliadins:conception and expression. Biochemical and Biophysical Research Communications,1997, 239:240-246
55. Eshel D, Miyara 1, Ailinng T, et al. pH regulates endoglucanase expression and virulence of Alternaria alternala in persimmon fruits. Molecular Plant-Microbe Interactions,2002,15:774-779
56. Feldman M, Lupton FGH, Millers TE. Wheats. In:Smartt J, Simmonds NW (eds) Evolution of crops. London:Longman,1995,184-192
57. Fernandez-Canon JM, Penalva MA. Molecular characterization of a gene encoding a homogentisate dioxygenase from Aspergillus nidulans and identification of its human and plant homologues. Journal of Biological Chemistry,1995,270:21199-21205
58. Ferraroni M, Seifert J, Travkin VM, et al. Crystal structure of the hydroxyquinol 1,2-dioxygenase from Nocardioides simplex 3E, a key enzyme involved in polychlorinated aromatics biodegradation. Journal of Biological Chemistry,2005, 280:21144-21154
59. Fido RJ, Bekes F, Gras PW, Tatham AS. Effects of α-, β-,γ-and ω-gliadins on the dough mixing properties of wheat flour. Journal of Cereal Science,1997,26:271-277
60. Francki M, Appels R. Wheat functional genomics and engineering crop improvement. Genome Biology,2002,3:1013.1-1013.5
61. Fu YX, Li WH. Statistical tests of neutrality of mutations. Genetics,1993, 133:693-709
62. Gaal AB, Neujahr HY. Maleylacetate reductase from Trichosporon cutaneum. Biochemical Journal,1980,185:783-786
63. Galili G. Heterologous expression of a wheat high molecular weight glutenin gene in Escherichia coli. Proc Natl Acad Sci USA,1989,86:7756-7760
64. Galka PWS, Ambrose SJ, Ross ARS, Abrams SR. Syntheses of deuterated jasmonates for mass spectrometry and metabolism studies. Journal of Labelled Compounds and Radiopharmaceuticals,2005,48:797-809
65. Ganzle MG, Loponen J, Gobbetti M. Proteolysis in sourdough fermentations: mechanisms and potential for improved bread quality. Trends in Food Science and Technology,2008,19:513-521
66. Gardiner DM, Osborne S, Kazan K, Manners JM. Low pH regulates the production of deoxynivalenol by Fusarium graminearum. Microbiology,2009,155:3149-3156
67. Garscha U, Jerneren F, Chung D, et al. Identification of dioxygenases required for Aspergillus development. Journal of Biological Chemistry,2007,282:34707-34718
68. Gellrich C, Schieberle P, Wieser H. Biochemical characterization and quantification of the storage protein (secalin) types in rye flour. Cereal Chemistry,2003,80:102-109
69. Gellrich C, Schieberle P, Wieser H. Biochemical characterization of y-75K secalins of rye Ⅱ. disulphide bonds. Cereal Chemistry,2004,81:296-299
70. Gellrich C, Schieberle P, Wieser H. Studies of partail amino acid sequences of γ-40K secalins of rye. Cereal Chemistry,2005,82:541-545
71. Gianibelli MC, Larroque OR, MacRitchie F, Wrigley CW. Biochemical, genetic, and molecular characterization of wheat endosperm proteins. American association of Cereal Chemists, Inc. Online review,2001,1-20
72. Gilbert J, Tekauz A. Review:Recent developments in research on fusarium head blight of wheat in Canada. Canadian Journal of Plant Pathology,2000,22:1-8
73. Giles RJ, Brown TA. GluDy allele variations in Aegilops tauschii and Triticum aestivum:implications for the origins of hexaploid wheats. Theoretical and Applied Genetics,2006,112:1563-1572
74. Glazebrook J, Chen W, Estes B, et al. Topology of the network integrating salicylate and jasmonate signal transduction derived from global expression phenotyping. Plant Journal,2003,34:217-228
75. Glazebrook J. Contrasting mechanisms of defense against biotrophic and necrotrophic pathogens. Annual Review of Phytopathology,2005,43:205-227
76. Gojobori T, Li WH, Graur D. Patterns of nucleotide substitution in pseudogenes and functional genes. Journal of Molecular Evolution,1982,18:360-369
77. Gorlach J, Volrath S, Knauf-Beiter G, et al. Benzothiadiazole, a novel class of inducers of systemic acquired resistance, activates gene expression and disease resistance in wheat. Plant Cell,1996,8:629-643
78. Goswami RS, Kistler HC. Heading for disaster:Fusarium graminearum on cereal crops. Molecular Plant Pathology,2004,5:515-525
79. Gu YQ, Crossman C, Kong XY, et al. Genomic organization of the complex a-gliadin gene loci in wheat. Theoretical and Applied Genetics,2004a,109:648-657
80. Gu YQ, Coleman-Derr D, Kong X, Anderson OD. Rapid genome evolution revealed by comparative sequence analysis of orthologous regions from four Triticeae genomes. Plant Physiology,2004b,135:459-470
81. Gu YQ, Wanjugi H, Coleman-Derr D, et al. Conserved globulin gene across eight grass genomes identify fundamental units of the loci encoding seed storage proteins. Funct Integr Genomics,2010,10:111-122
82. Guo ZF, Zhong M, Wei YM, et al. Characterization of two novel γ-gliadin genes encoded by K genome of Crithopsis delileana and evolution analysis with those from Triticeae. Genes & Genomics,2010,32:259-265
83. Hill WG, Robertson A. Linkage disequilibrium in finite populations. Theoretical and Applied Genetics,1968,38:226-231
84. Hisa CC, Anderson OD. Isolation and characterization of wheat w-gliadin genes. Theoretical and Applied Genetics,2001,103:37-44
85. Huang WE, Huang L, Preston GM, et al. Quantitative in situ assay of salicylic acid in tobacco leaves using a genetically modified biosensor strain of Acinetobacter sp. ADP1. Plant Journal,2006,46:1073-1083
86. Holzberg S, Brosio P, Gross C, et al. Barley stripe mosaic virus-induced gene silencing in a monocot plant. Plant Journal,2002,30:315-327
87. Hudson RR, Kaplan NL. Statistical properties of the number of recombination events in the history of a sample of DNA sequences. Genetics.1985,111:147-164
88. Hudson RR. Gene genealogies and the coalescent process. Oxford Surveys in Evolutionary Biology,1990,7:1-44
89. Ikeda TM, Nagamine T, Fukuoka H, Yano H. Identification of new Low-molecular-weight glutenin subunit genes in wheat. Theoretical and Applied Genetics,2002,104:680-687
90. Jansen C, von Wettstein D, Schafer W, et al. Infection patterns in barley and wheat spikes inoculated with wild-type and trichodiene synthase gene disrupted Fusarium graminearum. Proc Natl Acad Sci USA,2005,102:16892-16897
91. Ji L, Barrett T, Ayanbule O, et al. NCBI Peptidome:a new repository for mass spectrometry proteomics data. Nucleic Acids Research,2010,38:D731-D735
92. Kalin M, Neujahr HY, Weissmahr RN, et al. Phenol hydroxylase from Trichosporon cutaneum:gene cloning, sequence analysis, and functional expression in Escherichia coli. Journal of Bacteriology,1992,174:7112-7120
93. Kasarda DD, Bernardin JE, Nimmo CC. Wheat proteins in Advances in Cereal Science and Technology. Eds. Promeranz Y. St. Paul, MN. American Association of Cereal Chemists,1976a,1:158-236
94. Kasarda DD, Bernardin JE, Qualset CO. Relationship of gliadin components to chromosomes in hexaploid wheats(Triticum aestivum L.). Proc Natl Acad Sci USA, 1976b,73:3646-3650
95. Kasarda DD, Autran JC, Lew EJL, et al. N-terminal amino acid sequences of ω-gliadins and ω-secalins; implications for the evolution of prolamin genes. Biochimica et Biophysica Acta,1983,747:138-150
96. Kasarda DD, Okita TW, Bernardin JE, et al. Nucleic acid (cDNA) and amino acid sequences of a-type gliadins from wheat(Triticum aestivum). Proc Natl Acad Sci USA, 1984,81:4712-4716
97. Kasarda DD. Glutenin structure in relation to wheat quality. In wheat is unique: structure, composition, processing, end-use properties, and products. Edited by Y. Pomeranz. Minnesota:American Association of Cereal Chemists,1989,277-302
98. Kashkush K, Feldman M, Levy AA. Gene loss, silencing and activation in a newly synthesized wheat allotetraploid. Genetics,2002,160:1651-1659
99. Kawamichi H, Suzuki T. The cDNA-derived amino acid sequence of indoleamine dioxygenase like-myoglobin from the gastropod mollusc Omphalius pfeifferi. Journal of Protein Chemistry,1998,17:651-656
100. Kawano T. Roles of the reactive oxygen species-generating peroxidise reactions in plant defense and growth induction. Plant Cell Reports.2003,21:829-837
101. Kawaura K, Mochida K, Ogihara Y. Expression profile of two storage-protein gene families in hexaploid wheat revealed by large-scale analysis of expressed sequence tags. Plant Physiology,2005,139:1870-1880
102. Kelen M, Sanli N. Determination of pKa values of some auxins in methanol-water mixtures by reversed phase liquid chromatography and potentiometric methods. Journal of Brazilian Chemical Society,2009,20:133-140
103. Keller NP, Hohn TM. Metabolic pathway gene clusters in filamentous fungi. Fungal Genetics and Biology,1997,21:17-29
104. Kelly JK. A test of neutrality based on interlocus associations. Genetics,1997, 146:1197-1206
105. Khatkar BS, Fido RJ, Tatham AS, Schofield JD. Functional properties of wheat gliadins.1. effects on mixing characteristics and bread making quality. Journal of Cereal Science,2002a,35:299-306
106. Khatkar BS, Fido RJ, Tatham AS, Schofield JD. Functional properties of wheat gliadins. Ⅱ. effects on dynamic rheological properties of wheat gluten. Journal of Cereal Science,2002b,35:307-313
107. Klement Z, Goodman RN. The hepersensitive reaction to infection by bacterial plant pathogens. Annual Review of Phytopathology,1967,5:17-44
108. Kidd BN, Edgar CI, Kumar KK, et al. The mediator complex subunit PFT1 is a key regulator of jasmonate-dependent defense in Arabidopsis. Plant Cell,2009, 21:2237-2252
109. Kim JE, Lee HJ, Lee J, et al. Gibberella zeae chitin synthase genes, GzCHS5 and GzCHS7, are required for hyphal growth, perithecia formation, and pathogenicity. Current Genetics,2009,55:449-459
110. Koning F. The molecular basis of celiac disease. Journal of Molecular Recognition, 2003,16:333-336
111. Koornneef A, Pieterse CMJ. Cross talk in defense signalling. Plant Physiology,2008, 146,839-844
112. Kreis M, Forde BG, Rahman S, et al. Molecular evolution of the seed storage proteins. Journal of Molecular Biology,1985,183:499-502
113. Kruglyak L. The use of a genetic map of biallelic markers in linkage studies. Nature Genetics,1997,17:21-24
114. Kumar S, Tamura K, Nei M. MEGA 3:Integrated software for molecular evolutionary genetics analysis and sequence alignment. Brief in Bioinformatics,2004,5:150-163
115. Kupper U, Niedermann DM, Travaglini G, Lerch K. Isolation and characterization of the tyrosinase gene from Neurospora crassa. Journal of Biological Chemistry,1989, 264:17250-17258
116. Kurosawa E. Experimental studies on the nature of the substance secreted by the "bakanae" fungus. Nat Hist Soc Formosa,1926,16:213-227
117. Kwok S, Kellogg DE, McKinney N, et al. Effects of primer-template mismatches on the polymerase chain reaction:Human immunodeficiency virus type 1 model studies. Nucleic Acids Research,1990,18:999-1005
118. Laidman DL, Jones RG. Recent advances in the biochemistry of cereals. New York: Academic Press.1979.275-311
119. Laudert D, Weiler EW. Allene oxide synthase:a major control point in Arabidopsis thaliana octadecanoid signaling. Plant Journal,1998,15:675-684
120. Larkindale J, Knight MR. Protection against heat stress induced oxidative damage in Arabidopsis involves calcium, abscisic acid, ethylene, and salicylic acid. Plant Physiology,2002,128:682-695
121. Leon A, Rosell CM, de Barber CB. A differential scanning calorimetry study of wheat proteins. European Food Research and Technology,2003,217:13-16
122. Leon-Reyes A, Du Y, Koornneef A, et al. Ethylene signaling renders the jasmonate response of Arabidopsis insensitive to future suppression by salicylic acid. Molecular Plant-Microbe Interactions,2010,23:187-197
123. Lew EJL, Kuzmicky DD, Kasarda DD. Characterization of low-molecular-weight glutenin subunits by reversed-phase high-performance liquid chromatography, sodium dodecyl sulfate-polyacrylamide gel electrophoresis, and N-terminal amino-acid sequencing. Cereal Chemistry,1992,69:508-515
124. Li RX, Chen HB, Tu K, et al. Localized-statistical quantification of human serum proteome associated with type 2 diabetes. PLoS ONE,2008,3:e3224
125. Li G, Yen Y. Jasmonate and ethylene signaling pathway may mediate Fusarium head blight resistance in wheat. Crop Science,2008,48:1888-1896
126. Li GR, Ren ZL, Liu C, et al. Isolation and sequence analysis of a-gliadin genes from Dasypyrum breviaristatum. AtaTA Agronomica Sinica,2008,34:1097-1103
127. Li GR, Liu C, Zeng ZX, et al. Identification of a-gliadin genes in Dasypyrum in relation to evolution and breeding. Euphytica,2009,165:155-163
128. Li G, Zhang T, Wei P, et al. Sequence analysis of a-gliadin genes from Aegilops tauschii native to China. Asian Journal of Agricultural Sciences,2010a,2:128-135
129. Li G, Zhang T, Ban Y, Yang Z. Molecular characterization and evolutionary analysis of a-gliadin genes from Eremopyrum bonaepartis (Triticeae). Journal of Agricultural Science,2010b,2:30-36
130. Liu B, Vega JM, Feldman M. Rapid genomic changes in newly synthesized amphiploids of Triticum and Aegilops. II. Changes in low-copy coding DNA sequences. Genome,1998,41:535-542
131. Liu DC, Lan XJ, Yang ZJ, et al. A unique Aegilops tauschii genotype needless to immature embryo culture in cross with wheat. Acta Botanica Sinica,2002,44:708-713
132. Liu DC, Fang H. Study on the role of Chinese Ae. tauschii in the evolution of Chinese common wheat landraces. Southwest China Journal of Agricultural Sciences,2003, 16:32-35
133. Long H, Wei YM, Yan ZH, et al. Classification of wheat low-molecular-weight glutenin subunit genes and its chromosome assignment by developing group-specific primers. Theoretical and Applied Genetics,2005,111:1251-1259
134. Lopez MA, Bannenberg G, Castresana C. Controlling hormone signaling is a plant and pathogen challenge for growth and survival. Current Opinion in Plant Biology,2008, 11:420-427
135. Ma LJ, van der Does HC, Borkovich KA, et al. Comparative genomics reveals mobile pathogenicity chromosomes in Fusarium. Nature,2010,464:367-373
136. Ma ZC, Wei YM, Yan ZH, et al. Characterization of a-gliadin genes from diploid wheats and the comparative analysis with those from polyploid wheats. Russian Journal of Genetics,2007,43:1286-1293
137. Macnicol PK, Jacobsen JV. Endosperm acidification and related metabolic changes in the developing barley grain. Plant Physiology,1992,98:1098-1104
138. Makandar R, Essig JS, Schapaugh MA, et al. Genetically engineered resistance to Fusarium head blight in wheat by expression of Arabidopsis NPR1. Molecular Plant-Microbe Interactions,2006,19:123-129
139. Makandar R, Nalam R, Shah J. Interaction between salicylic acid and jasmonic acid signaling in Arabidopsis and wheat impacts macroconidia germination of the Fusarium head blight fungus, Fusarium graminearum. In:Proceedings of the 9th International Conference on Arabidopsis Research, Montreal, Canada, July 23-27, 2008,148
140. Makandar R, Nalam V, Chaturvedi R, et al. Involvement of salicylate and jasmonate signaling pathways in Arabidopsis interaction with Fusarium graminearum. Molecular Plant-Microbe Interactions,2010,23:861-870
141. Malamy J, Sanchez-Casas P, Hennig J, et al. Dissection of the salicylic acid signaling pathway in Tobacco. Molecular Plant-Microbe Interactions,1996,9:474-482
142. Malonek S, Bomke C, Bornberg-Bauer E, et al. Distribution of gibberellin biosynthetic genes and gibberellin production in the Gibberella fujikuroi species complex. Phytochemistry,2005,66:1296-1311
143. Martin-Udiroz M, Madrid MP, Roncero MI. Role of chitin synthase genes in Fusarium oxysporum. Microbiology,2004,150:3175-3187
144. Maruyama N, Ichise K, Katsube T, et al. Identification of major wheat allergens by means of the Escherichia coli expression system. European Journal of Biochemistry, 1998,255:739-745
145. Masci S, Lew EJL, Lafiandra D, et al. Characterization of low molecular weight glutenin subunits in durum wheat by reversed-phase high-performance liquid chromatography and N-terminal sequencing. Cereal Chemistry,1995,72:100-104
146. Masci S, D'Ovidio R, Lafiandra D, Kasarda DD. Characterization of a low-molecular-weight glutenin subunit gene from bread and corresponding protein that represents a major subunit of the glutenin polymer. Plant Physiology,1998, 118:1147-1158
147. Masci S, Rovelli L, Kasarda DD, et al. Characterization and chromosomal localization of C-type low-molecular-weight glutenin subunits in the bread wheat cultivar Chinese Spring. Theoretical and Applied Genetics,2002,104:422-428
148. Maucher H, Hause B, Feussner I, et al. Allene oxide synthases of barley (Hordeum vulgare cv. Salome):tissue specific regulation in seedling development. Plant Journal, 2000,21:199-213
149. Mei C, Qi M, Sheng G, Yang Y. Inducible overexpression of a rice allene oxide synthase gene increases the endogenous jasmonic acid level, PR gene expression, and host resistance to fungal infection. Molecular Plant-Microbe Interactions,2006, 19:1127-1137
150. McAdam SN, Sollid LM. Getting to grips with gluten. Gut 2000,47:743
151. McCartney CA, Somers DJ, Fedak G, Cao W. Haplotype diversity at fusarium head blight resistance QTLs in wheat. Theoretical and Applied Genetics,2004, 109:261-271
152. McCorkle GM, Yeh WK. Fletcher P. Ornston LN. Repetitions in the NH2-terminal amino acid sequence of beta-ketoadipate enol-lactone hydrolase from Pseudomonas putida. Journal of Biological Chemistry,1980,255:6335-6341
153. McFadden ES, Sears ER. The origin of Triticum spelta and its free-threshing hexaploid relatives. Journal of Heredity,1946,37:81-89,107-116
154. McManus R, Kelleher D. Celiac disease-the villain unmasked? New England Journal of Medicine,2003,348:2573-2574
155. Mesterhazy A. Types and components of resistance to Fusarium head blight of wheat. Plant Breeding,1995,114:377-386
156. Metakovsky EV, Branlard G. Genetic diversity of French common wheat germplasm based on gliadin alleles. Theoretical and Applied Genetics,1998,96:209-218
157. Mika A, Boenisch MJ, Hopff D, Luthje S. Membrane-bound guaiacol peroxidases from maize (Zea mays L.) roots are regulated by methyl jasmonate, salicylic acid, and pathogen elicitors. Journal of Experimental Botany,2010,61:831-841
158. Millar AA, Jacobsen JV, Ross JJ, et al. Seed dormancy and ABA metabolism in Arabidopsis and barley:the role of ABA 8'-hydroxylase. Plant Journal,2006, 45:942-954
159. Miller D, Blackwell BA. Biosynthesis of 3-acetyldeoxynivalenol and other metabolites by Fusarium culmorum HLK 1503 in a stirred jar fermentor. Canadian Journal of Botany,1986.64:1-5
160. Miller VM, Xia H, Marrs GL, et al. Allele-specific silencing of dominant disease genes. Proc Natl Acad Sci USA,2003,100:7195-7200
161. Molberg 0, Uhlen AK, Jensen T, et al. Mapping of gluten T cell epitopes in the bread wheat ancestors:implications for celiac disease. Gastroenterology,2005,128:393-401
162. Molina A, Gorlach J, Volrath S, Ryals J. Wheat genes encoding two types of PR-1 proteins are pathogen inducible, but do not respond to activators of systemic acquired resistance. Molecular Plant-Microbe Interactions,1999,12:53-58
163. Muller SW, Wieser H. The location of disulphide bonds in a-type gliadins. Journal of Cereal Science,1995,22:21-27
164. Miiller S, Wieser H, Popineau Y. Disulphide bonds of γ46-gliadin. Journal of Cereal Science,1998,27:23-25
165. Murray EE, Lotzer J, Eberle M. Codon usage in plant genes. Nucleic Acids research, 1989,17:477-498
166. Murray FR, Skerritt JH, Appels R. A gene from the Sec2(Gli-R2) locus of a wheat 2RS.2BL chromosomal translocation line. Theoretical and Applied Genetics,2001, 102:431-439
167. Nagy IJ, Takacs I, Juhasz A, et al. Identification of a new class of recombinant prolamin genes in wheat. Genome,2005,48:840-847
168. Navarro L, Bari R, Achard P, et al. DELLAs control plant immune responses by modulating the balance of jasmonic acid and salicylic acid signaling. Current Biology, 2008,18:650-655
169. Nei M. Molecular Evolutionary Genetics. New York:Columbia University Press 1987
170. Neujahr HY, Lindsjo S, Varga JM. Oxidation of phenols by cells and cell-free enzymes from Candida tropicalis. Antonie van Leeuwenhoek,1974,40:209-216
171. Nevo E, Korol AB, Beiles A, Fahima T. Evolution of Wild Emmer and wheat Improvement:population genetics, genetic resources, and genome organization of wheat's progenitor, Triticum dicoccoides. Springer-Verlag Berlin Heidelberg New York,2002.1-17
172. Ohan N, Heikkila JJ. Involvement of differential gene expression and mRNA stability in the developmental regulation of the hsp 30 gene family in heat-shocked Xenopus laevis embryos. Developmental Genetics,1995,17:176-184
173. Okita TW, Cheesbrough V, Reeves CD. Evolution and heterogeneity of the α-/β-type and γ-type gliadin DNA sequences. Journal of Biological Chemistry,1985, 260:8203-8213
174. Ornebro J, Nylander T, Eliasson AC, et al. Adsorption of the high molecular weight glutenin subunit 1Dx5 compared to the 58-kDa central repetitive domain and α-gliadins. Journal of Cereal Science,2001,34:141-150
175. Ornebro J, Nylander T, Eliasson AC, et al. The behavior of the high molecular-weight glutenin subunit 1DX5, the 58kDa central repetitive domain and α-gliadins at the air-aqueous interface. Journal of Cereal Science,2003,38:147-156
176. Osborne TB. The protein of the wheat kernel. Washington, D.C.:Carnegie Institute, 1907, Publication No.84
177. Ozkan H, Levy AA, Feldman M. Allopolyploidy-induced rapid genome evolution in the wheat(Aegilops-Triticum) group. Plant cell,2001,13:1735-1747
178. Paabo S, Irwin DM, Wilson AC. DNA damage promotes jumping between templates during enzymatic amplification. Journal of Biological Chemistry,1990, 265:4718-4721
179. Panozzo C, Nawara M, Suski C, et al. Aerobic and anaerobic NAD+ metabolism in Saccharomyces cerevisiae. FEBS Letters,2002,517:97-102
180. Paulitz TC. Fusarium head blight:a re-emerging disease. Phytoprotection,1999, 80:127-133
181. Payne PI, Holt LM, Jackson EA, Law CN. Wheat storage proteins:their genetics and potential for manipulation by plant breeding. Philos Trans R Soc London Ser B 1984a, 304:359-371
182. Payne PI, Jackson EA, Holt LM, Law CN. Genetic linkage between endosperm storage protein genes on each of the short arms of chromosomes 1A and 1B in wheat. Theoretical and Applid Genetics,1984b,67:235-243
183. Payne PI. Genetics of wheat storage proteins and the effect of allelic variation on bread-making quality. Annual Review of Plant Physiology,1987,38:141-153
184. Payne PI, Law CN, Mudd EE. Control by homoeologous group 1 chromosomes of the high-molecular-weight subunits of glutenin, a major protein of wheat endosperm. Theoretical and Applid Genetics,1980,58:113-120
185. Pel HJ, de Winde JH, Archer DB, et al. Genome sequencing and analysis of the versatile cell factory Aspergillus niger CBS 513.88. Nature Biotechnology,2007, 25:221-231
186. Penninckx IAMA, Thomma BPHJ, Buchala A, et al. Concomitant activation of jasmonate and ethylene response pathways is required for induction of a plant defensin gene in Arabidopsis. Plant Cell,1998,10:2103-2113
187. Pieterse CM, van Loon LC. NPR1:the spider in the web of induced resistance signaling pathways. Current Opinion in Plant Biology,2004,7:456-464
188. Pieterse CMJ, Leon-Reyes A, Van der Ent S, et al. Networking by small-molecule hormones in plant immunity. Nature Chemical Biology,2009,5:308-316
189. Piston F, Dorado G, Martin A, Barro F. Cloning of nine γ-gliadin mRNAs (cDNAs) from wheat and the molecular characterization of comparative transcript levels of y-gliadin subclasses. Journal of Cereal Science,2006,43:120-128
190. Pollitt S, Zalkin H. Role of primary structure and disulfide bond formation in β-lactamase secretion. Journal of Bacteriology,1983,153:27-32
191. Popineau Y, Pineau F. Investigation of surface hydrophobicities of purified gliadins by hydrophobic interaction chromatography. reverse-phage high performance chromatography and a polar ligand binding. Journal of Cereal Science,1987, 5:215-231
192. Press WH, Teukolsky SA, Vetterling WT, Flannery BP. Numerical recipes in C. The art of Scientific Computing. Cambridge:Cambridge University Press 1992
193. Prusky D, McEvoy JL, Leverentz B, Conway WS. Local modulation of host pH by Colletotrichum species as a mechanism to increase virulence. Molecular Plant-Microbe Interactions,2001,14:1105-13
194. Qi PF, Wei YM, Yue YW, et al. Biochemical and molecular characterization of gliadins. Molecular Biology,2006a,40:713-723
195. Qi PF, Yue YW, Wei YM, et al. Molecular characterization of α-gliadin genes from wild emmer wheat (Triticum dicoccoides). DNA sequence,2006b,17:415-421
196. Qi PF, Wei YM, Ouellet T, et al. The y-gliadin multigene family in common wheat (Triticum aestivum) and its closely related species. BMC Genomics,2009,10:168
197. Qi PF, Wei YM, Chen Q, et al. Identification of novel a-gliadin genes. Genome, 2011a,54:244-252
198. Qi PF, Wei YM, Chen GY, et al. Development of chromosome 6D-specific markers for a-gliadin genes and their use in assessing dynamic changes at the Gli-2 loci. Molecular Breeding,2011b, doi:10.1007/sl 1032-010-9539-5
199. Qin X, Zeevaart JAD. The 9-cis-epoxycarotenoid cleavage reaction is the key regulatory step of abscisic acid biosynthesis in water-stressed bean. Proc Natl Acad Sci USA,1999,96:15354-15361
200. Rafalski JA, Scheets K, Metzler M, et al. Developmentally regulated plant genes:the nucleotide sequence of a wheat gliadin genomic clone. EMBO Journal,1984, 3:1409-1415
201. Rafalski J A. Structure of wheat gamma-gliadin genes. Gene,1986,43:221-229
202. Reeves CD, Okita TW. Analysis of α/β-type gliadin genes from diploid and hexaploid wheats. Gene,1987,52:257-266
203. Ren J, Wen L, Gao X, et al. DOG1.0:Illustrator of protein domain structures. Cell Research,2009,19:271-273
204. Robert-Seilaniantz A, Navarro L, Bari R, et al. Pathological hormone imbalances. Current Opinion in Plant Biology,2007,10:372-379
205. Rocheleau HJ, Zheng W, Gulden S, et al. Comparative gene expression profiling of major plant hormone pathways during infection by Fusarium graminearum. In: Ouellet, T., Leger, D. (Eds), Proceedings of the Sixth Canadian Workshop on Fusarium Head Blight, Ottawa, ON, Canada,2009, pp.103, http://www.cwfhb.org /pastprograms.html.
206. Romanova YA, Gubareva NK, Konarev AV, et al. Analysis of gliadin polymorphism in a Triricum spelta L. collection. Russian Journal of Genetics,2001,37:1054-1060
207. Ross ARS, Ambrose SJ, Cutler AJ, et al. Determination of endogenous and supplied deuterated abscisic acid in plant tissues by high performance liquid chromatography-electrospray ionization tandem mass spectrometry with multiple reaction monitoring. Analytical Biochemistry,2004,329:324-333
208. Rozas J, Sanchez-Del Barrio JC, Messeguer X, Rozas R. DnaSP, DNA polymorphism analyses by the coalescentand other methods. Bioinformatics,2003,19:2496-2497
209. Rungis D, Hamberger B, Berube Y, et al. Efficient genetic mapping of single nucleotide polymorphisms based upon DNA mismatch digestion. Molecular Breeding, 2005,16:261-270
210. Sabelli P, Shewry PR. Characterization and organization of gene families at the Gli-1 loci of bread and durum wheats by restriction fragment analysis. Theoretical and Applied Genetics,1991,83:209-216
211. Salcedo G, Prada J, Aragoncillo C. Low-molecular-weight gliadin-like proteins from wheat endosperm. Phytochemistry,1979,18:725-727
212. Salcedo G, Prada J, Sanchez-Monge R, Aragoncillo C. Aneuploid analysis of low-molecular-weight gliadins from wheat. Theoretical and Applied Genetics,1980, 56:65-69
213. Santha R, Savithri HS, Rao NA, Vaidyanathan CS.2,3-Dihydroxybenzoic acid decarboxylase from Aspergillus niger. A novel decarboxylase. European Journal of Biochemistry,1995,230:104-110
214. Savitch LV, Subramaniam R, Allard GC, et al. The GLK1'regulon'encodes disease defense related proteins and confers resistance to Fuasarium graminearum in Arabidopsis. Biochemical and Biophysical Research Communications,2007, 359:234-238
215. Scheets K, Hedgcoth C. Nucleotide sequence of a gamma gliadin gene:Comparisons with other gamma gliadin sequences show the structure of gamma gliadin genes and the general primary structure of gamma gliadins. Plant Science,1988,57:141-150
216. Schofield JD, Bottomley RC, Timms MF, Booth MR. The effect of heat on wheat gluten and the involvement of sulphydryl-disulphide interchange reactions. Journal of Cereal Science,1983,1:241-253
217. Scofield SR, Huang L, Brandt AS, et al. Development of a virus-induced gene-silencing system for hexaploid wheat and its use in functional analysis of the Lr21-mediated leaf rust resistance pathway. Plant Physiology,2005,138:2165-2173
218. Shan L, Molberg (?), Parrot I, et al. Structural basis for gluten intolerance in celiac disease. Science,2002,297:2275-2279
219. Shewry PR, Halford NG, Belton PS, Tatham AS. The structure and properties of gluten: an elastic protein from wheat grain. Phil Trans Roy Soc Lond B Biol Sci,2002, 357:133-142
220. Shewry PR, Napier JA, Tatham AS. Seed storage proteins:structures and biosynthesis. Plant Cell,1995,7:945-956
221. Shewry PR, Popineau Y, Lafiandra D, Belton P. Wheat glutenin subunits and dough elastricity:findings of the EURO-WHEAT project. Trends in Food Science and Technology,2001,11:433-441
222. Shewry PR, Tatham AS, Lazzeri P. Biotechnology of wheat quality. Journal of the Science of Food and Agriculture,1997,73:397-406
223. Shewry PR, Halford NG. Cereal seed storage proteins:structures, properties and role in grain utilization. Journal of Experimental Botany,2002,53:947-958
224. Shewry PR, Tatham AS. The prolamin storage proteins of cereal seeds:structure and evolution. Biochemical Journal,1990,267:1-12
225. Shim WB, Woloshuk CP. Regulation of fumonisin B(1) biosynthesis and conidiation in Fusarium verticillioides by a cyclin-like (C-type) gene, FCC1. Applied and Environmental Microbiology,2001.67:1607-1612
226. Sinha RC, Savard ME, Lau R. Production of monoclonal antibodies for the specific detection of deoxynivalenol and 15-acetyldeoxynivalenol by ELISA. The Journal of Agricultural and Food Chemistry,1995,43:1740-1744
227. Somers DJ, Fedak G, Savard M. Molecular mapping of novel genes controlling Fusarium head blight resistance and deoxynivalenol accumulation in spring wheat. Genome,2003,46:555-564
228. Spaenij-Dekking L, Kooy-Winkelaar Y, van Veelen P, et al. Natural variation in toxicity of wheat:potential for selection of nontoxic varieties for celiac disease patients. Gastroenterology,2005,129:797-806
229. Spoel SH, Dong X. Making sense of hormone crosstalk during plant immune responses. Cell Host & Microbe,2008,12:348-351
230. Sugiyama T, Rafalski A, Soell D. The Nucleotide sequence of a wheat gamma-gliadin genomic clone. Plant Science,1986,44:205-209
231. Sumner-Smith M, Rafalski JA, Sugiyama T, et al. Conservation and variability of wheat α/β-gliadin genes. Nucleic Acids Research,1985,13:3905-3916
232. Tajima F. Statistical method for testing the neutral mutation hypothesis by DNA polymorphism. Genetics,1989,123:585-595
233. Tang QY, Feng MG. DPS Data processing system:Experimental design, statistical analysis, and data mining. Beijing:Science Press,2007
234. Tao HP, Cornell DG, Kasarda DD. Surface and optical properties of wheat glutenin monolayers. Journal of Cereal Science,1989,10:5-18
235. Tao HP, Kasarda DD. Two-dimensional gel sequencing of LMW-glutenin subunits. Journal of Experimental Botany,1989,40:1015-1020
236. Tatham AS, Shewry PR. The S-poor prolamins of wheat, barley and rye. Journal of Cereal Science,1995,22:1-16
237. Taylor IB, Burbidge A, Thompson AJ. Control of abscisic acid synthesis. Journal of Experimental Botany,2000,51:1563-1574
238. Thaler JS, Owen B, Higgins VJ. The role of the jasmonate response in plant susceptibility to diverse pathogens with a range of lifestyles. Plant Physiology,2004, 135:53-538
239. Thatcher LF, Manners JM, Kazan K. Fusarium oxysporum hijacks COIl-mediated jasmonate signaling to promote disease development in Arabidopsis. Plant Journal, 2009,58:927-939
240. Thomma BPHJ, Penninckx IAMA, Broekaert WF, Cammue BPA. The complexity of disease signaling in Arabidopsis. Current Opinion in Immunology,2001,13:63-68
241. Thompson JD, Gibson TJ, Plewniak F, et al. The CLUSTAL X windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research,1997,25:4876-4882
242. Ton J, Flors V. Mauch-Mani B. The multifaceted role of ABA in disease resistance. Trends in Plant Science,2009,14:310-317
243. Tsavkelova EA, Klimova SY, Cherdyntseva TA, et al. Microbial producers of plant growth stimulators and their practical use:a review. Applied Biochemistry and Microbiology,2006,42:117-126
244. Tsavkelova EA, Bomke C, Netrusov Al, et al. Production of gibberellic acids by an orchid-associated Fusarium proliferatum strain. Fungal Genetics and Biology,2008, 45:1393-1403
245. Uknes S, Mauch-Mani B, Moyer M, et al. Acquired resistance in Arabidopsis. Plant Cell,1992,4:645-656
246. Vaccino P, Becker H, Brandolini A, Salamini F, Kilian B. A catalogue of Triticum monococcum genes encoding toxic and immunogenic peptides for celiac disease patients. Mol Genet Genomics,2009,281:289-300
247. van Herpen TWJM, Goryunova SV, van der Schoot J, et al. Alpha-gliadin genes from the A, B and D genomes of wheat contain different sets of celiac disease epitopes. BMC Genomics,2006,7:1
248. Varga JM, Neujahr HY. Purification and properties of catechol-1,2-oxygenase from Trichosporon cutaneum. European Journal of Biochemistry,1970,12:427-434
249. Viswanathan VK, Krcmarik K, Cianciotto NP. Template secondary structure promotes polymerase jumping during PCR amplification. Biotechniques,1999,27:508-511
250. von Buren M, Luthy J, Hiibner P. A spelt-specific y-glaidin gene:discovery and detection. Theoretical and Applied Genetics,2000,100:271-279
251. Wang JR, Wang L, Gulden S, et al. RNA profiling of fusarium head blight-resistant wheat addition lines containing the Thinopyrum elongtum chromosome 7E. Canadian Journal of Plant Pathology,2010,32:188-214
252. Walter S, Nicholson P, Doohan FM. Action and reaction of host and pathogen during Fusarium head blight disease. New Phytologist,2010,185:54-66
253. Wang R R-C, von Bothmer R, Dvorak J, et al. Genome symbols in the Triticeae (Poaceae). In:Wang R R-C, Jensen KB, Jaussi C (eds). Proceedings of the 2nd international Triticeae Symposium, Logan, Utah, USA.1996,29-34
254. Ward ER, Uknes SJ, Williams SC, et al. Coordinate gene activity in response to agents that induce systemic acquired resistance. Plant Cell,1991,3:1085-1094
255. Weegels PL, Marseille JP, Bosveld P, Hamer RJ. Large-scale separation of gliadins and their bread-making quality. Journal of Cereal Science,1994,20:253-264
256. Wicker T, Guyot R, Yahiaoui N, Keller B. CACTA transposons in Triticeae, A diverse family of high-copy repetitive elements. Plant Physiology,2003a,132:52-63
257. Wicker T, Yahiaoui N, Guyot R, et al. Rapid genome divergence at orthologous low molecular weight glutenin loci of the A and Am genomes of wheat. Plant Cell,2003b, 15:1186-1197
258. Wieser H. Comparative investigations of gluten proteins from different wheat species. I. Qualitative and quantitative composition of gluten protein types. European Food Research and Technology,2000,211:262-268
259. Wieser H. Comparative investigations of gluten proteins from different wheat species. Ⅲ. N-terminal amino acid sequences of a-gliadins potentially toxic for celiac patients. European Food Research and Technology,2001,213:183-186
260. Wieser H, Belitz HD, Idar D, Ashkenazi A. Coeliac activity of the gliadin peptides CT-1 and CT-2. Z Lebensm Unters Forsch,1986,182:115-117
261. Wieser H, Springer G, Belitz HD. Toxicity of different wheat gliadins in coeliac disease. Z Lebensm Forsch,1982,175:321-326
262. Windsor ML, Miborrow BV, McFarlane IJ. The uptake of (+)-S-and (-)-R-abscisic acid by suspension culture cells of Hopbush (Dodonaea viscose). Plant Physiology, 1992,100:54-62
263. Woychik JH, Boundy JA, Dimler RJ. Starch gel-electrophoresis of wheat gluten proteins with concentrated urea. Archives of Biochemistry and Biophysics,1961, 94:477-482
264. Wright JD. Fungal degradation of benzoic acid and related compounds. World Journal of Microbiology and Biotechnology,1993,9:9-16
265. Xie Z, Wang C, Wang K, Wang S, Li X, Zhang Z, Ma W, Yan Y. Molecular characterization of the celiac disease epitope domains in α-gliadin genes in Aegilops tauschii and hexaploid wheats (Triticum aestivum L.). Theoretical and Applied Genetics,2010.121:1239-1251
266. Xu JH, Messing J. Amplification of prolamin storage protein genes in different subfamilies of the Poaceae. Theoretical and Applied Genetics,2009,119:1397-1412
267. Xu XM, Berrie AM. Epidemiology of mycotoxigenic fungi associated with Fusarium ear blight and apple blue mould:a review. Food Additives and Contaminants,2005, 22:290-301
268. Xu YB, Li HP, Zhang JB, et al. Disruption of the chitin synthase gene CHS1 from Fusarium asiaticum results in an altered structure of cell walls and reduced virulence. Fungal Genetics and Biology,2010,47:205-215
269. Yen C, Yang JL. The biosystematics study on the Triticeae tribe.Ⅰ. Triticum-Aegilops complex. Beijing:China Agricultural Press,1999
270. You IS, Ghosal D, Gunsalus IC. Nucleotide sequence analysis of the Pseudomonas putida PpG7 salicylate hydroxylase gene (nahG) and its 3'-flanking region. Biochemistry,1991,12:1635-1641
271. Yu GY, Muehlbauer GJ. Benzothiadiazole-induced gene expression in wheat spikes does not provide resistance to Fusarium head blight. Physiological and Molecular Plant Pathology,2001,59:129-136
272. Zaharia LI, Galka MM, Ambrose SJ, et al. Preparation of deuterated abscisic acid metabolites for use in mass spectrometry and feeding studies. Journal of Labelled Cmpounds and Radiopharmaceuticals,2005,48:435-445
273. Zhang LQ, Liu DC, Yan ZH, et al. Rapid changes of microsatellite flanking sequence in the allopolyploidization of new synthesized hexaploid wheat. Science in China Ser. C,2004,47:553-561
274. Zheng YL, Zhou YH, Wei YM. The Applied Research on Molecular Biology of Wheat. Chengdu:Sichuan Science and Technology Press,2001
275.郭天财.小麦及其亲源的分类。河南农业大学学报,1985,19：208-229
276. 姜玉英,曾娟,周益林等.小麦病虫害发生于监控。北京：中国农业出版社,2008,1-39
277.李振崎.商鸿生,魏宁生.麦类病害。北京：中国农业出版社,1997,1-19
278.王爱丽,裴玉贺,张倩,晏月明.小麦非醇溶蛋白的研究进展。首都师范大学学报(自然科学版),2006,27：68-74
279.王玉海,代彦涛,郑美等.小麦-山羊草复合群的起源与进化研究进展。中国农学通报,2010,26：1]6-118
280.余遥,饶世达,任正隆等.四川小麦。成都：四川科学技术出版社,1998,103-160,391-428
281.喻章,马奇祥,王成俊.小麦病虫害及其防治。成都：四川大学出版社,2002,36-40
282.翟凤林,李宗智,王光瑞等.作物品质育种。北京：农业出版社,1991,3-67,207-249
283.张效梅,白建荣,贾旭.乌拉尔图小麦的遗传学研究进展。西北植物学报,2004,24：1972-1978