利用特异小麦资源创制育种新材料及其遗传评价
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摘要
小麦是世界上最广泛种植的作物,它是世界上占35%人口的主要粮食作物。但普通小麦品种遗传基础狭窄,遗传差异小,制约其产量和品质的进一步提高。为了提高栽培小麦的产量和抗性,提高其遗传背景是非常重要的。在小麦属及其近缘属物种中,具有许多改良小麦所需的目标基因,它们能有效的丰富小麦的遗传变异和拓宽其遗传多样性。在向普通小麦转移外源遗传物质的过程中,远缘杂交和合成双二倍体扮演了重要的角色。矮秆波兰小麦(Triticum turgidum ssp.polonicum L.,AABB,2n=4x=28,Dwarfing Polish Wheat,DPW)是颜济、杨俊良教授在进行联合国粮农组织小麦植物资源考察项目时,采集于新疆吐鲁番的一份新的四倍体小麦矮秆资源,株高约68cm,属矮秆群,是迄今为止波兰小麦唯一的矮秆变异材料。它对赤霉酸反应不敏感,具有长穗、多小穗、多分蘖、裸粒等特征。华山新麦草(Psathyrostachys huashanica Kengex Guo)是禾本科(Poaceae)小麦族(Triticeae)新麦草属(Psathyrostachys Nevski)的一个多年生二倍体(2n=14)物种,含Ns基因组。它是我国特有的禾草植物,仅生长在陕西华山,与同属的其他物种有较大的形态差异和形成间断地理分布。华山新麦草具有抗寒、抗旱、耐瘠薄、早熟、优质、高抗小麦条锈病和全蚀病,中抗赤霉病等特点。由于华山新麦草独特的分类学地位、特殊的地理分布和作为小麦族的重要基因资源,目前该物种已被列为我国珍稀濒危一级保护植物和急需保护的农作物近缘种。
本文围绕这两个特异种质材料进行相关研究。分析了矮秆波兰小麦矮秆基因的遗传特点和构建了第一张四倍体小麦矮秆基因的SSR分子图谱;在不使用幼胚培养等辅助条件下,在大田自然环境中,通过属间杂交和秋水仙碱加倍合成了矮秆波兰小麦与节节麦的双二倍体(SHW-DPW);并利用质体乙酰-CoA羧化酶(ACCase)基因(Acc-1)研究了SHW-DPW与新疆稻麦(Triticum petropavlovskyi Udacz.et Migusch.)的系统发育关系及其新疆稻麦可能的起源途径;研究比较了phKL基因与人工Ph基因突变系在小麦-华山新麦草杂种间的可杂交性与诱导部分同源染色体配对的作用大小;利用秋水仙碱处理普通小麦与华山新麦草的杂种F_1植株,第一次人工合成了小麦属与新麦草属间的双二倍体,并对它的形态学、分子细胞遗传学、抗病性等进行了研究。主要研究结果如下:
1.矮秆波兰小麦是一份特异的来自中国新疆的具有矮秆性状的波兰小麦,株高约68cm。遗传学分析表明矮秆波兰小麦所携带的矮秆性状是由一对不完全显性基因控制,暂定名为Rht-dp。为了标记这个四倍体小麦的矮秆基因,利用矮秆波兰小麦与高秆波兰小麦(AS304)杂交,建立了F_2分离群体。共选用153对均匀分布于小麦A、B染色体的SSR引物,在供试亲本间扩增,其中43对引物在双亲之同表现差异,约占所用引物的28.1%。采用BSA方法,将已筛选的43对引物在矮、高池间进行扩增,发现引物WMC511、Xgwm495、Xgwm113和Xgwm251在矮、高池间表现多态,表明这些引物与Rht-dp连锁,并将该矮秆基因初步定位到4BS上。进一步用这四个标记对F_2代124个单株进行PCR扩增,根据扩增结果,采用Mapmaker 3.0软件计算遗传距离,结果显示,Rht-dp位于WMC511和Xgwm495、Xgwm113、Xgwm251之间,它们的排列顺序可能是WMC511—Rht-dp—Xgwm495—Xgwm113—Xgwm251,它们之间的遗传距离分别为0.4 cM、4.1 cM、2.5 cM、10.3 cM。研究结果为标记辅助育种提供了分子选择工具,同时也为进一步精细定位和图位克隆Rht-dp基因奠定了基础。
2.未经幼胚拯救及激素处理,第一次成功获得了矮秆波兰小麦与节节麦(AS60和AS65)的F_1杂种。矮秆波兰小麦与节节麦(AS60和AS65)两个杂交组合的杂交结实率分别为1.67%和0.60%。只有T.turgidum ssp.polonicum(DPW)×Ae.tauschii(AS60)组合发芽良好,而且获得了24株杂种植株。所有F_1杂种植株生长旺盛,形态特征与面包小麦相似。F_1植株具有一些明显的来自于亲本矮秆波兰小麦和节节麦的特征,且完全不育。杂种F_1花粉母细胞减数分裂MI染色体主要以单价体形式出现,平均每个细胞有20.56个单价体,0.22个二价体,且全为棒状,未观察到环状二价体和多价体。此外,F_1杂种的成功获得对研究普通小麦的起源和育种利用具有重要意义。
3.通过秋水仙碱处理矮秆波兰小麦与节节麦(AS60)的F_1杂种植株,第一次人工合成了一个六倍体小麦材料(SHW-DPW)。两株F_1植株的自交结实率分别为8.27%和15.31%。SHW-DPW的形态特征与F_1植株相似,但生长更旺盛。所有SHW-DPW植株形态相似,而且表达了亲本矮秆波兰小麦与节节麦的一些特征。SHW-DPW植株的颖片相对较软及穗轴节很短,这与其他合成六倍体小麦是不同的。SHW-DPW的穗子形态与新疆稻麦非常相似。SHW-DPW是可育的,2n=42条染色体的植株的自交结实率为58.95%,非整倍体的自交结实率为45.63%。在花粉母细胞减数分裂中期Ⅰ,新合成小麦SHW-DPW的42条染色体的配对构型为0.43个单价体、20.77个二价体和0.01个三价体,说明新双二倍体在细胞遗传上是相对稳定的。并对SHW-DPW在研究新疆稻麦起源和小麦育种利用方面的潜在价值进行了讨论。
4.利用质体乙酰-CoA羧化酶(ACCase)基因(Acc-1)分析了我们合成的六倍体小麦SHW-DPW与新疆稻麦(Triticum petropavlovskyi Udacz.et Migusch.)及其小麦属(Triticum)和山羊草属(Aegilops)共47个物种的分子系统与进化关系。系统发育分析使用最大似然法(ML)和MJ构建系统发育树。所有新疆稻麦与波兰小麦材料Acc-1基因序列发生了大量的核苷酸替换,具有很高的遗传多样性。在新疆稻麦A、B和D基因组Acc-1基因序列上,以A基因组的遗传多样性最高,D基因组最低,B基因组居中。SHW-DPW的Acc-1基因序列分别与其母本矮秆波兰小麦和节节麦聚在一起,与新疆稻麦聚在不同的分支,结果表明SHW-DPW与新疆稻麦的亲缘关系比较远。在MJ网状结构中,新疆稻麦A、B基因组Acc-1基因序列首先与新疆的波兰小麦和普通小麦聚在一起,具有最高的序列相似性,揭示了他们之间特定的祖先—后代关系。在ML和MJ系统发育分析中,新疆稻麦D基因组Acc-1基因序列与普通小麦单独聚在一个分支,而5份节节麦与SHW-DPW聚在另一个分支,说明新疆稻麦与普通小麦的亲缘关系要近于新疆稻麦与节节麦的亲缘关系。以上研究结果表明新疆稻麦起源于独立的异源多倍化事件或由普通小麦经过简单的突变形成的可能性很小,它更可能是波兰小麦与普通小麦之间通过自发的基因渗入或自然驯化而产生。
5.在普通小麦地方品种开县罗汉麦(Triticum aestivum L.,2n=6x=42,AABBDD)中天然存在促进小麦-外源杂种部分同源染色体配对的基因phKL。本研究在没有采用幼胚培养技术条件下,利用华山新麦草和普通小麦直接杂交,成功获得了杂种F_1。研究比较了phKL基因与人工Ph基因突变系在小麦-华山新麦草杂种间的可杂交性与诱导部分同源染色体配对的作用大小。在所有杂交组合中,杂种开县罗汉麦×华山新麦草的杂交结实率最高,为3.18%。在花粉母细胞减数分裂MI,开县罗汉麦×华山新麦草杂种的染色体配对构型为21.70个单价体、2.68个棒状二价体、0.34个环状二价体、0.06个三价体和0.02个四价体。与人工Ph基因突变系相比,开县罗汉麦与华山新麦草杂种的部分同源染色体配对水平显著增加,表现在棒状二价体、环状二价体和三价体的数量变多而单价体数量减少。研究结果表明,phKL在诱导小麦-华山新麦草部分同源染色体配对的作用能力比Ph1和Ph2强。因此,在转移亲源关系较远的外源遗传物质时,含有phKL基因的开县罗汉麦可能是一个更好的供体材料。
6.在向普通小麦转移外源遗传物质的过程中,远缘杂交和合成双二倍体扮演了重要的角色。利用秋水仙碱处理普通小麦与华山新麦草的杂种F_1植株,第一次人工合成了小麦属与新麦草属间的双二倍体(AABBDDNsNs,PHW-SA)。PHW-SA的形态特征与杂种F_1植株相似,且生长显得更旺盛。所有PHW-SA植株形态相似,亲本J-11与华山新麦草的一些形态特征在新双二倍体中得到了完全表达。例如,PHW-SA植株节间呈紫色和基部小穗上有绒毛,这些都是来自华山新麦草的特点。PHW-SA是完全可育的,其自交结实率为37.10%-77.38%,平均59.11%。通过根尖细胞有丝分裂观察,PHW-SA的染色体数目变化范围2n=51到2n=56,70.59%的植株具有2n=56条染色体。在花粉母细胞减数分裂MI,PHW-SA(2n=56)染色体配对构型为1.15个单价体、27.34个二价体、0.03个三价体和0.02个四价体。在观察的1226个花粉母细胞中,在MI 49.79%染色体完全配对,表明该双二倍体(2n=56)在细胞遗传上是相对稳定的。在非整倍体植株中,减数分裂后期Ⅰ和Ⅱ出现数目不等的落后染色体和微核。本研究报道了普通小麦与华山新麦草的双二倍体的形成、形态和细胞学特征,并对该双二倍体在理论及应用方面的特殊价值进行了讨论。
7.为了评估华山新麦草遗传物质在小麦育种中潜在的利用价值,本研究采用高分子量谷蛋白亚基分析(SDS-PAGE)、醇溶蛋白分析(A-PAGE)、Giemsa-C带、基因组原位杂交(GISH)和随机扩增多态性DNA(RAPD)方法对八倍体PHW-SA的分子细胞遗传学特点进行了分析,并检测了它的抗条锈病能力。Giemsa-C带分析表明,华山新麦草染色体具有强烈的端带,并在PHW-SA中完全表达。用华山新麦草的基因组DNA作探针,对PHW-SA进行基因组原位杂交分析,双二倍体56条染色体由10条完整的华山新麦草染色体、36条小麦染色体和10条小麦—华山新麦草易位染色体组成。种子贮藏蛋白分析表明PHW-SA表达了华山新麦草特异的条带,且出现了亲本物种没有的新带纹或亲本物种的带纹消失。PHW-SA的抗条锈病检测表明来自华山新麦草的抗条锈性在双二倍体中完全表达。选用5个对华山新麦草N_S基因组特异的RAPD引物,对双二倍体及其亲本进行扩增,结果表明能够在双二倍体和两个亲本中扩增出相同的带纹。对PHW-SA可以作为向普通小麦中转移有益基因的一种新的遗传资源进行了讨论。
Bread wheat(Triticum aestivum L.) is the most widely grown cereal in the world.It is the staple food for 35%of the world's population,and is becoming increasingly important in the developing world.To meet the demands for high yielding and stress-resistant wheat cultivars,it is desirable to increase the genetic base of this crop. Many wild species of the tribe Triticeae(Poaceae) are valuable sources for resistance to diseases,salinity,drought and insect pests.Wide crosses and synthetic amphiploids have played an important role in introgressing desirable traits from related species into cultivated wheat.Dwarfing polish wheat(Triticum turgidurn ssp.polonicum,DPW) is a tertraploid species(2n = 4x = 28,AABB),and was collected from Tulufan,Xinjiang, China.It is the only one dwarfing accession of T.turgidurn ssp.polonicum,with the height of about 68 cm,which is insensitive in gibberellin acid reaction,and is characterized as dwarfing,more tillers,more length of spike,more spikelets per spike and length of glume.Psathyrostachys huashanica Keng ex,Guo(2n=2x=14,NsNs) is a perennial species in the genus Psathyrostachys Nevski,and is located only in Hua Shan, Shanxi province,China.P.huashanica has been served as an important resource to wheat improvement because of its disease resistance,resistant ability to unfavorable environment,dwarfing and tremendous genetic diversity.At present,the amount of the population is much less than the past and being at the edge of extinction.Owing to distributing limitation and importance as breeding material for germplasm storage,it was listed into the first class of national protected rare plants and imperatively protected wild species in relation to crops.
In the present study,we studied the genetic character and molecular mapping of dwarfing gene of dwarfing polish wheat using SSR.By colchicine treatment of the hybrid plants between dwarfing polish wheat and Aegilops tauschii Cosson.,a new synthetic hexaploid wheat(SHW-DPW) was artificially obtained for the fist time.The relationship between SHW-DPW and Triticum petropavlovskyi Udacz.et Miguch.and the origin of T. petropavlovskyi were discussed based on the sequences of a singly-copy nuclear gene encoding plastid acetyl-CoA carboxylase(Acc-1).The effects of phKL gene on crossability and homoeologous pairing of T.aestivum×p.huashanica hybrids were comparatively analyzed.The hybrid plants between T.aestivum and P.huashanica were treated with colchicine,a new intergeneric amphiploid(PHW-SA) was obtained for the fist time,and the morphology,fertility,molecular cytogenetic characterization and disease responses of the amphiploid were reported.The results were as follows:
1.Dwarfing polish wheat(DPW) is a dwarfing accession of T.turgidum ssp. polonicum from Xinjiang of China.The mean plant height of DPW is 68cm.Genetic analysis of F_2、BC_1 and BC_2 populations derived from DPW×T.turgidum ssp.polonicum (AS304) indicated that the culm length in DPW was determined by a single partially dominant gene,and which was named as Rht-dp.In order to map the height-reducing gene Rht-dp,a total of 153 SSR primer pairs covering A and B wheat chromosomes were used for polymorphism survey of DPW and AS304.Among these primers,43(28.1%) showed polymorphism between the parents.By Bulk Segregant Analysis(BSA) of dwarfing and tall pools,four pairs of primer located on chromosome arm 4BS,WMC511、Xgwm495、Xgwm 113 and Xgwm251,were found being linked to Rht-dp.The 124 F_2 individuals were then amplified with the four markers.Further molecular mapping showed that Rht-dp was linked with SSR markers in the order of WMC511-Rht-dp-Xgwm495-Xgwm113-Xgwm251 with genetic distances of 0.4cM、4.1cM、2.5cM and 10.3cM using the software Mapmaker 3.0,respectively.The molecular markers developed in this study will be useful for marker-assisted selection of alternative height-reducing genes,and to better map and clone Rht-dp gene.
2.The artificial hybridization between dwarfing polish wheat and two accessions of Ae.tauschii(AS60 and AS65) was carried out,and the F_1 hybrids were obtained successfully without using embryo rescue techniques for the first time.The crossabilities of hybrids T.turgidum ssp.polonicum(DPW)×Ae.tauschii(AS60) and T.turgidum ssp. polonicurn(DPW)×Ae.tauschii(AS65) were 1.67%and 0.60%respectively.Only the hybrids of T.turgidum ssp.polonicum(DPW)×Ae.tauschii(AS60) germinated well,and 24 F_1 hybrid plants were obtained.All the F_1 hybrid plants grew vigorously,and the morphological traits were similar to bread wheat.The F_1 plants had some obvious traits inherited from the parents and were completely sterile.Chromosome pairing in the hybrid was characterized by a large number of univalents,with an average of 20.56 and 0.22 bivalents per PMC,and no ring bivalents and multivalents were observed.Furthermore,the potential value of the F_1 hybrids between T.turgidum ssp.polonicurn and Ae.tauschii for studying wheat origin and breeding are discussed.
3.By colchicine treatment of the hybrid plants between T.turgidum ssp.polonicum (DPW) from Xinjiang in China and Ae.tauschii from Middle East,a new synthetic hexaploid wheat(SHW-DPW) was artificially obtained for the fist time.The seed sets of the two F_1 plants were 8.27%and 15.31%,respectively.The morphology of the SHW-DPW plants was comparable to that of the primary F_1 plants,and the SHW-DPW plants appeared more robust.All the SHW-DPW plants were uniform in morphology and had some obvious traits inherited from T.turgidurn ssp.polonicurn(DPW) and Ae.tauschii. Especially,the glumes were very soft and the rachis intemodia was short,which was different from other synthetic hexaploid wheat.Their spikes were morphologically similar to those of T.petropavlovskyi.The SHW-DPW plants were fertile,with 58.95%selfed seed set for the euhexaploids and 45.63%for the aneuploids,respectively.The meiosis of the SHW-DPW plants was quite regular,which showed a pairing configuration of 0.43 univalents,20.77 bivalents and 0.01 trivalents.The rate of chromosome association of the aneuploids was comparatively lower than that in the euploid individuals.The potential utilization for wheat improvement and study for the origin of T.petropavlovskyi are discussed in the present paper.
4.Based on the sequences of a singly-copy nuclear gene encoding plastid acetyl-CoA carboxylase(Acc-1),a total of 47 accessions Triticum and Aegilops representing diploid, tetraploid and hexaploid were used to estimate the origin of T.petropavlovskyi and the relationship of synthetic hexaploid wheat(SHW-DPW) and T.petropavlovskyi. Phylogenetic analysis was performed using maximum likelihood(ML),and median-joining (MJ) networks.Genetic diversity of all T.petropavlovskyi and T.turgidum ssp. polonicum accessions were high owing to a number of nucleotide substitutions among the wheat species.Among the three genomes,the A and B genome Acc-1 gene of T. petropavlovskyi have the more polymorphisms than D genome.The Acc-1 gene of SHW-DPW was consistently more closely related to the parent T.polonicum and Ae. tauschii than to T.petropavlovskyi,it showed that the relationship of SHW-DPW and T. petropavlovskyi was distant.Furthermore,it suggested that the A and B genome haplotypes from Acc-1 loci show that T.petropavlovskyi shares the highest average sequence identity with T.turgidum ssp.polonicum from Xinjiang and T.aestivum,and reveals specific progenitor-descendant relationships in the MJ networks based on intron sequence.In the ML tree of intron + sy data sets,the D genome sequences of the Acc-1 genes from T. petropavlovskyi are identical to the sequences of the D genome orthologs in T.aestivum, while the relationship of T.petropavlovskyi and Ae.tauschii are most distant,which is in agreement with the result of MJ networks analysis.Findings of this study reduced the probability of an independent allopolyploidization event and a single mutation in T. aestivum in the origin of T.petropavlovskyi and indicated a greater degree of gene flow between T.aestivum and T.turgidum ssp.polonicum leading to T.petropavlovskyi.It is most likely that T.petropavlovskyi was introduced from T.turgidum ssp.polonicum to T. aestivum via a spontaneous introgression or breeding effort.
5.In the natural populations of common wheat landrace Kaixianluohanmai,there was a phKL gene which promotes homoeologous pairing of wheat-alien hybrids.In the present study,the effects of phKL gene on crossability and homoeologous pairing of T.aestivum×P.huashanica hybrids were comparatively analyzed.The crossability of the hybrid between Sichuan wheat landrace Kaixianluohanmai and P.huashanica was highest in all the hybrid combinations with 3.18%.The hybrids T.aestivum(Kaixianluohanmai)×p. huashanica showed a pairing configuration of 21.70 univalents + 2.68 rod bivalents + 0.34 ring bivalents + 0.06 trivalents + 0.02 quadrivalents and 3.54 chiasma per PMC at MI. However,the chiasma in hybrids of CS,CSphlb,CSph2a and CSph2b with P.huashanica was 0.56,1.90,0.87 and 0.60 respectively.Compared with the hybrids of CS,CSphlb, CSph2a and CSph2b with P.huashanica,a significant increase in the chiasma of homoeologous pairing was observed in the hybrids of T.aestivum(Kaixianluohanrnai)×p. huashanica.The effects were shown in the increment of rod bivalents,ring bivalents and trivalents and reduction of univalents.The results indicated that phKL showed a higher effect on promoting homoeologous pairing than phl and ph2 in T.aestivum×p. huashanica.The wheat landrace Kaixianluohanmai with phKL gene is probably a more desirable material for alien gene transferring than Ph2-deficiency lines.
6.Wide crosses and synthetic amphiploids have played an important role in introgressing desirable traits from related species into cultivated wheat.The hybrid plants between T.aestivum cv.J-11 and P.huashanica were treated with colchicine,a new intergeneric amphiploid(PHW-SA) was obtained for the fist time.The morphological characteristics of PHW-SA resembled the parent J-11.The PHW-SA plants have purple internode and floss in basal spikelet,which are inherited from P.huashanica parent.The somatic chromosome number varied from 2n=51 to 2n=56,with 70.59%plants having 56 chromosomes.At metaphaseⅠ,PHW-SA(2n=56) showed average of 1.15 univalents, 27.34 bivalents,0.03 trivalents and 0.02 tetravalents per cell,and complete chromosome pairing was found in 49.79%of the PMCs.Lagging chromosomes and micronuclei were observed in the aneuploid cells.A survey of diseases resistance revealed that the stripe rust resistance from P.huashanica was totally expressed and powdery mildew was suppressed in the PHW-SA.The fertility of PHW-SA was 37.10%-77.38%,with average of 59.11%. PHW-SA can be used as a donor source in wheat breeding programme.
7.A new wheat-P,huashanica amphiploid,PHW-SA,was characterized by molecular cytological observation and tested for stripe rust resistance in order to evaluate the potential use of the P.huashanica for wheat improvement.The PHW-SA plants were deeply covered with floss in basal spikelet and purple internode,which is characteristic of the P.huashanica parent.Feulgen staining of the somatic metaphases revealed that the chromosome number varied from 51 to 56.At metaphaseⅠ,PHW-SA(2n=56) regularly showed averagely 1.15 univalents and 27.34(24-28) bivalents per cell,and complete chromosome pairing was found in 49.79%of the pollen mother cells(PMCs),indicating a degree of cytological stability.Giemsa-C banding showed that the P.huashanica chromosomes in PHW-SA produced strong bands only in their terminal regions.Genomic in situ hybridization(GISH) analysis suggested that PHW-SA carried 10 entire P. huashanica chromosomes,36 wheat chromosomes and 10 wheat-P,huashanica translocated chromosomes.Seeds storage proteins separated by acid polyacrylamide gel electrophoresis(A-PAGE) and sodium dodecyl sulphate-polyacrylarnide gel electrophoresis(SDS-PAGE) showed that PHW-SA expressed some of P.huashanica specific gliadin and glutenin bands,and some new bands appeared.A survey of diseases resistance revealed that the stripe rust resistance from P.huashanica was totally expressed and powdery mildew was suppressed in the PHW-SA.Random amplified polymorphic DNA(RAPD) analysis verified the presence of DNA of the parents in PHW-SA.The present study showed that the amphiploid PHW-SA could serve as novel sources for transfer of valuable genes to wheat.
本文围绕这两个特异种质材料进行相关研究。分析了矮秆波兰小麦矮秆基因的遗传特点和构建了第一张四倍体小麦矮秆基因的SSR分子图谱;在不使用幼胚培养等辅助条件下,在大田自然环境中,通过属间杂交和秋水仙碱加倍合成了矮秆波兰小麦与节节麦的双二倍体(SHW-DPW);并利用质体乙酰-CoA羧化酶(ACCase)基因(Acc-1)研究了SHW-DPW与新疆稻麦(Triticum petropavlovskyi Udacz.et Migusch.)的系统发育关系及其新疆稻麦可能的起源途径;研究比较了phKL基因与人工Ph基因突变系在小麦-华山新麦草杂种间的可杂交性与诱导部分同源染色体配对的作用大小;利用秋水仙碱处理普通小麦与华山新麦草的杂种F_1植株,第一次人工合成了小麦属与新麦草属间的双二倍体,并对它的形态学、分子细胞遗传学、抗病性等进行了研究。主要研究结果如下:
1.矮秆波兰小麦是一份特异的来自中国新疆的具有矮秆性状的波兰小麦,株高约68cm。遗传学分析表明矮秆波兰小麦所携带的矮秆性状是由一对不完全显性基因控制,暂定名为Rht-dp。为了标记这个四倍体小麦的矮秆基因,利用矮秆波兰小麦与高秆波兰小麦(AS304)杂交,建立了F_2分离群体。共选用153对均匀分布于小麦A、B染色体的SSR引物,在供试亲本间扩增,其中43对引物在双亲之同表现差异,约占所用引物的28.1%。采用BSA方法,将已筛选的43对引物在矮、高池间进行扩增,发现引物WMC511、Xgwm495、Xgwm113和Xgwm251在矮、高池间表现多态,表明这些引物与Rht-dp连锁,并将该矮秆基因初步定位到4BS上。进一步用这四个标记对F_2代124个单株进行PCR扩增,根据扩增结果,采用Mapmaker 3.0软件计算遗传距离,结果显示,Rht-dp位于WMC511和Xgwm495、Xgwm113、Xgwm251之间,它们的排列顺序可能是WMC511—Rht-dp—Xgwm495—Xgwm113—Xgwm251,它们之间的遗传距离分别为0.4 cM、4.1 cM、2.5 cM、10.3 cM。研究结果为标记辅助育种提供了分子选择工具,同时也为进一步精细定位和图位克隆Rht-dp基因奠定了基础。
2.未经幼胚拯救及激素处理,第一次成功获得了矮秆波兰小麦与节节麦(AS60和AS65)的F_1杂种。矮秆波兰小麦与节节麦(AS60和AS65)两个杂交组合的杂交结实率分别为1.67%和0.60%。只有T.turgidum ssp.polonicum(DPW)×Ae.tauschii(AS60)组合发芽良好,而且获得了24株杂种植株。所有F_1杂种植株生长旺盛,形态特征与面包小麦相似。F_1植株具有一些明显的来自于亲本矮秆波兰小麦和节节麦的特征,且完全不育。杂种F_1花粉母细胞减数分裂MI染色体主要以单价体形式出现,平均每个细胞有20.56个单价体,0.22个二价体,且全为棒状,未观察到环状二价体和多价体。此外,F_1杂种的成功获得对研究普通小麦的起源和育种利用具有重要意义。
3.通过秋水仙碱处理矮秆波兰小麦与节节麦(AS60)的F_1杂种植株,第一次人工合成了一个六倍体小麦材料(SHW-DPW)。两株F_1植株的自交结实率分别为8.27%和15.31%。SHW-DPW的形态特征与F_1植株相似,但生长更旺盛。所有SHW-DPW植株形态相似,而且表达了亲本矮秆波兰小麦与节节麦的一些特征。SHW-DPW植株的颖片相对较软及穗轴节很短,这与其他合成六倍体小麦是不同的。SHW-DPW的穗子形态与新疆稻麦非常相似。SHW-DPW是可育的,2n=42条染色体的植株的自交结实率为58.95%,非整倍体的自交结实率为45.63%。在花粉母细胞减数分裂中期Ⅰ,新合成小麦SHW-DPW的42条染色体的配对构型为0.43个单价体、20.77个二价体和0.01个三价体,说明新双二倍体在细胞遗传上是相对稳定的。并对SHW-DPW在研究新疆稻麦起源和小麦育种利用方面的潜在价值进行了讨论。
4.利用质体乙酰-CoA羧化酶(ACCase)基因(Acc-1)分析了我们合成的六倍体小麦SHW-DPW与新疆稻麦(Triticum petropavlovskyi Udacz.et Migusch.)及其小麦属(Triticum)和山羊草属(Aegilops)共47个物种的分子系统与进化关系。系统发育分析使用最大似然法(ML)和MJ构建系统发育树。所有新疆稻麦与波兰小麦材料Acc-1基因序列发生了大量的核苷酸替换,具有很高的遗传多样性。在新疆稻麦A、B和D基因组Acc-1基因序列上,以A基因组的遗传多样性最高,D基因组最低,B基因组居中。SHW-DPW的Acc-1基因序列分别与其母本矮秆波兰小麦和节节麦聚在一起,与新疆稻麦聚在不同的分支,结果表明SHW-DPW与新疆稻麦的亲缘关系比较远。在MJ网状结构中,新疆稻麦A、B基因组Acc-1基因序列首先与新疆的波兰小麦和普通小麦聚在一起,具有最高的序列相似性,揭示了他们之间特定的祖先—后代关系。在ML和MJ系统发育分析中,新疆稻麦D基因组Acc-1基因序列与普通小麦单独聚在一个分支,而5份节节麦与SHW-DPW聚在另一个分支,说明新疆稻麦与普通小麦的亲缘关系要近于新疆稻麦与节节麦的亲缘关系。以上研究结果表明新疆稻麦起源于独立的异源多倍化事件或由普通小麦经过简单的突变形成的可能性很小,它更可能是波兰小麦与普通小麦之间通过自发的基因渗入或自然驯化而产生。
5.在普通小麦地方品种开县罗汉麦(Triticum aestivum L.,2n=6x=42,AABBDD)中天然存在促进小麦-外源杂种部分同源染色体配对的基因phKL。本研究在没有采用幼胚培养技术条件下,利用华山新麦草和普通小麦直接杂交,成功获得了杂种F_1。研究比较了phKL基因与人工Ph基因突变系在小麦-华山新麦草杂种间的可杂交性与诱导部分同源染色体配对的作用大小。在所有杂交组合中,杂种开县罗汉麦×华山新麦草的杂交结实率最高,为3.18%。在花粉母细胞减数分裂MI,开县罗汉麦×华山新麦草杂种的染色体配对构型为21.70个单价体、2.68个棒状二价体、0.34个环状二价体、0.06个三价体和0.02个四价体。与人工Ph基因突变系相比,开县罗汉麦与华山新麦草杂种的部分同源染色体配对水平显著增加,表现在棒状二价体、环状二价体和三价体的数量变多而单价体数量减少。研究结果表明,phKL在诱导小麦-华山新麦草部分同源染色体配对的作用能力比Ph1和Ph2强。因此,在转移亲源关系较远的外源遗传物质时,含有phKL基因的开县罗汉麦可能是一个更好的供体材料。
6.在向普通小麦转移外源遗传物质的过程中,远缘杂交和合成双二倍体扮演了重要的角色。利用秋水仙碱处理普通小麦与华山新麦草的杂种F_1植株,第一次人工合成了小麦属与新麦草属间的双二倍体(AABBDDNsNs,PHW-SA)。PHW-SA的形态特征与杂种F_1植株相似,且生长显得更旺盛。所有PHW-SA植株形态相似,亲本J-11与华山新麦草的一些形态特征在新双二倍体中得到了完全表达。例如,PHW-SA植株节间呈紫色和基部小穗上有绒毛,这些都是来自华山新麦草的特点。PHW-SA是完全可育的,其自交结实率为37.10%-77.38%,平均59.11%。通过根尖细胞有丝分裂观察,PHW-SA的染色体数目变化范围2n=51到2n=56,70.59%的植株具有2n=56条染色体。在花粉母细胞减数分裂MI,PHW-SA(2n=56)染色体配对构型为1.15个单价体、27.34个二价体、0.03个三价体和0.02个四价体。在观察的1226个花粉母细胞中,在MI 49.79%染色体完全配对,表明该双二倍体(2n=56)在细胞遗传上是相对稳定的。在非整倍体植株中,减数分裂后期Ⅰ和Ⅱ出现数目不等的落后染色体和微核。本研究报道了普通小麦与华山新麦草的双二倍体的形成、形态和细胞学特征,并对该双二倍体在理论及应用方面的特殊价值进行了讨论。
7.为了评估华山新麦草遗传物质在小麦育种中潜在的利用价值,本研究采用高分子量谷蛋白亚基分析(SDS-PAGE)、醇溶蛋白分析(A-PAGE)、Giemsa-C带、基因组原位杂交(GISH)和随机扩增多态性DNA(RAPD)方法对八倍体PHW-SA的分子细胞遗传学特点进行了分析,并检测了它的抗条锈病能力。Giemsa-C带分析表明,华山新麦草染色体具有强烈的端带,并在PHW-SA中完全表达。用华山新麦草的基因组DNA作探针,对PHW-SA进行基因组原位杂交分析,双二倍体56条染色体由10条完整的华山新麦草染色体、36条小麦染色体和10条小麦—华山新麦草易位染色体组成。种子贮藏蛋白分析表明PHW-SA表达了华山新麦草特异的条带,且出现了亲本物种没有的新带纹或亲本物种的带纹消失。PHW-SA的抗条锈病检测表明来自华山新麦草的抗条锈性在双二倍体中完全表达。选用5个对华山新麦草N_S基因组特异的RAPD引物,对双二倍体及其亲本进行扩增,结果表明能够在双二倍体和两个亲本中扩增出相同的带纹。对PHW-SA可以作为向普通小麦中转移有益基因的一种新的遗传资源进行了讨论。
Bread wheat(Triticum aestivum L.) is the most widely grown cereal in the world.It is the staple food for 35%of the world's population,and is becoming increasingly important in the developing world.To meet the demands for high yielding and stress-resistant wheat cultivars,it is desirable to increase the genetic base of this crop. Many wild species of the tribe Triticeae(Poaceae) are valuable sources for resistance to diseases,salinity,drought and insect pests.Wide crosses and synthetic amphiploids have played an important role in introgressing desirable traits from related species into cultivated wheat.Dwarfing polish wheat(Triticum turgidurn ssp.polonicum,DPW) is a tertraploid species(2n = 4x = 28,AABB),and was collected from Tulufan,Xinjiang, China.It is the only one dwarfing accession of T.turgidurn ssp.polonicum,with the height of about 68 cm,which is insensitive in gibberellin acid reaction,and is characterized as dwarfing,more tillers,more length of spike,more spikelets per spike and length of glume.Psathyrostachys huashanica Keng ex,Guo(2n=2x=14,NsNs) is a perennial species in the genus Psathyrostachys Nevski,and is located only in Hua Shan, Shanxi province,China.P.huashanica has been served as an important resource to wheat improvement because of its disease resistance,resistant ability to unfavorable environment,dwarfing and tremendous genetic diversity.At present,the amount of the population is much less than the past and being at the edge of extinction.Owing to distributing limitation and importance as breeding material for germplasm storage,it was listed into the first class of national protected rare plants and imperatively protected wild species in relation to crops.
In the present study,we studied the genetic character and molecular mapping of dwarfing gene of dwarfing polish wheat using SSR.By colchicine treatment of the hybrid plants between dwarfing polish wheat and Aegilops tauschii Cosson.,a new synthetic hexaploid wheat(SHW-DPW) was artificially obtained for the fist time.The relationship between SHW-DPW and Triticum petropavlovskyi Udacz.et Miguch.and the origin of T. petropavlovskyi were discussed based on the sequences of a singly-copy nuclear gene encoding plastid acetyl-CoA carboxylase(Acc-1).The effects of phKL gene on crossability and homoeologous pairing of T.aestivum×p.huashanica hybrids were comparatively analyzed.The hybrid plants between T.aestivum and P.huashanica were treated with colchicine,a new intergeneric amphiploid(PHW-SA) was obtained for the fist time,and the morphology,fertility,molecular cytogenetic characterization and disease responses of the amphiploid were reported.The results were as follows:
1.Dwarfing polish wheat(DPW) is a dwarfing accession of T.turgidum ssp. polonicum from Xinjiang of China.The mean plant height of DPW is 68cm.Genetic analysis of F_2、BC_1 and BC_2 populations derived from DPW×T.turgidum ssp.polonicum (AS304) indicated that the culm length in DPW was determined by a single partially dominant gene,and which was named as Rht-dp.In order to map the height-reducing gene Rht-dp,a total of 153 SSR primer pairs covering A and B wheat chromosomes were used for polymorphism survey of DPW and AS304.Among these primers,43(28.1%) showed polymorphism between the parents.By Bulk Segregant Analysis(BSA) of dwarfing and tall pools,four pairs of primer located on chromosome arm 4BS,WMC511、Xgwm495、Xgwm 113 and Xgwm251,were found being linked to Rht-dp.The 124 F_2 individuals were then amplified with the four markers.Further molecular mapping showed that Rht-dp was linked with SSR markers in the order of WMC511-Rht-dp-Xgwm495-Xgwm113-Xgwm251 with genetic distances of 0.4cM、4.1cM、2.5cM and 10.3cM using the software Mapmaker 3.0,respectively.The molecular markers developed in this study will be useful for marker-assisted selection of alternative height-reducing genes,and to better map and clone Rht-dp gene.
2.The artificial hybridization between dwarfing polish wheat and two accessions of Ae.tauschii(AS60 and AS65) was carried out,and the F_1 hybrids were obtained successfully without using embryo rescue techniques for the first time.The crossabilities of hybrids T.turgidum ssp.polonicum(DPW)×Ae.tauschii(AS60) and T.turgidum ssp. polonicurn(DPW)×Ae.tauschii(AS65) were 1.67%and 0.60%respectively.Only the hybrids of T.turgidum ssp.polonicum(DPW)×Ae.tauschii(AS60) germinated well,and 24 F_1 hybrid plants were obtained.All the F_1 hybrid plants grew vigorously,and the morphological traits were similar to bread wheat.The F_1 plants had some obvious traits inherited from the parents and were completely sterile.Chromosome pairing in the hybrid was characterized by a large number of univalents,with an average of 20.56 and 0.22 bivalents per PMC,and no ring bivalents and multivalents were observed.Furthermore,the potential value of the F_1 hybrids between T.turgidum ssp.polonicurn and Ae.tauschii for studying wheat origin and breeding are discussed.
3.By colchicine treatment of the hybrid plants between T.turgidum ssp.polonicum (DPW) from Xinjiang in China and Ae.tauschii from Middle East,a new synthetic hexaploid wheat(SHW-DPW) was artificially obtained for the fist time.The seed sets of the two F_1 plants were 8.27%and 15.31%,respectively.The morphology of the SHW-DPW plants was comparable to that of the primary F_1 plants,and the SHW-DPW plants appeared more robust.All the SHW-DPW plants were uniform in morphology and had some obvious traits inherited from T.turgidurn ssp.polonicurn(DPW) and Ae.tauschii. Especially,the glumes were very soft and the rachis intemodia was short,which was different from other synthetic hexaploid wheat.Their spikes were morphologically similar to those of T.petropavlovskyi.The SHW-DPW plants were fertile,with 58.95%selfed seed set for the euhexaploids and 45.63%for the aneuploids,respectively.The meiosis of the SHW-DPW plants was quite regular,which showed a pairing configuration of 0.43 univalents,20.77 bivalents and 0.01 trivalents.The rate of chromosome association of the aneuploids was comparatively lower than that in the euploid individuals.The potential utilization for wheat improvement and study for the origin of T.petropavlovskyi are discussed in the present paper.
4.Based on the sequences of a singly-copy nuclear gene encoding plastid acetyl-CoA carboxylase(Acc-1),a total of 47 accessions Triticum and Aegilops representing diploid, tetraploid and hexaploid were used to estimate the origin of T.petropavlovskyi and the relationship of synthetic hexaploid wheat(SHW-DPW) and T.petropavlovskyi. Phylogenetic analysis was performed using maximum likelihood(ML),and median-joining (MJ) networks.Genetic diversity of all T.petropavlovskyi and T.turgidum ssp. polonicum accessions were high owing to a number of nucleotide substitutions among the wheat species.Among the three genomes,the A and B genome Acc-1 gene of T. petropavlovskyi have the more polymorphisms than D genome.The Acc-1 gene of SHW-DPW was consistently more closely related to the parent T.polonicum and Ae. tauschii than to T.petropavlovskyi,it showed that the relationship of SHW-DPW and T. petropavlovskyi was distant.Furthermore,it suggested that the A and B genome haplotypes from Acc-1 loci show that T.petropavlovskyi shares the highest average sequence identity with T.turgidum ssp.polonicum from Xinjiang and T.aestivum,and reveals specific progenitor-descendant relationships in the MJ networks based on intron sequence.In the ML tree of intron + sy data sets,the D genome sequences of the Acc-1 genes from T. petropavlovskyi are identical to the sequences of the D genome orthologs in T.aestivum, while the relationship of T.petropavlovskyi and Ae.tauschii are most distant,which is in agreement with the result of MJ networks analysis.Findings of this study reduced the probability of an independent allopolyploidization event and a single mutation in T. aestivum in the origin of T.petropavlovskyi and indicated a greater degree of gene flow between T.aestivum and T.turgidum ssp.polonicum leading to T.petropavlovskyi.It is most likely that T.petropavlovskyi was introduced from T.turgidum ssp.polonicum to T. aestivum via a spontaneous introgression or breeding effort.
5.In the natural populations of common wheat landrace Kaixianluohanmai,there was a phKL gene which promotes homoeologous pairing of wheat-alien hybrids.In the present study,the effects of phKL gene on crossability and homoeologous pairing of T.aestivum×P.huashanica hybrids were comparatively analyzed.The crossability of the hybrid between Sichuan wheat landrace Kaixianluohanmai and P.huashanica was highest in all the hybrid combinations with 3.18%.The hybrids T.aestivum(Kaixianluohanmai)×p. huashanica showed a pairing configuration of 21.70 univalents + 2.68 rod bivalents + 0.34 ring bivalents + 0.06 trivalents + 0.02 quadrivalents and 3.54 chiasma per PMC at MI. However,the chiasma in hybrids of CS,CSphlb,CSph2a and CSph2b with P.huashanica was 0.56,1.90,0.87 and 0.60 respectively.Compared with the hybrids of CS,CSphlb, CSph2a and CSph2b with P.huashanica,a significant increase in the chiasma of homoeologous pairing was observed in the hybrids of T.aestivum(Kaixianluohanrnai)×p. huashanica.The effects were shown in the increment of rod bivalents,ring bivalents and trivalents and reduction of univalents.The results indicated that phKL showed a higher effect on promoting homoeologous pairing than phl and ph2 in T.aestivum×p. huashanica.The wheat landrace Kaixianluohanmai with phKL gene is probably a more desirable material for alien gene transferring than Ph2-deficiency lines.
6.Wide crosses and synthetic amphiploids have played an important role in introgressing desirable traits from related species into cultivated wheat.The hybrid plants between T.aestivum cv.J-11 and P.huashanica were treated with colchicine,a new intergeneric amphiploid(PHW-SA) was obtained for the fist time.The morphological characteristics of PHW-SA resembled the parent J-11.The PHW-SA plants have purple internode and floss in basal spikelet,which are inherited from P.huashanica parent.The somatic chromosome number varied from 2n=51 to 2n=56,with 70.59%plants having 56 chromosomes.At metaphaseⅠ,PHW-SA(2n=56) showed average of 1.15 univalents, 27.34 bivalents,0.03 trivalents and 0.02 tetravalents per cell,and complete chromosome pairing was found in 49.79%of the PMCs.Lagging chromosomes and micronuclei were observed in the aneuploid cells.A survey of diseases resistance revealed that the stripe rust resistance from P.huashanica was totally expressed and powdery mildew was suppressed in the PHW-SA.The fertility of PHW-SA was 37.10%-77.38%,with average of 59.11%. PHW-SA can be used as a donor source in wheat breeding programme.
7.A new wheat-P,huashanica amphiploid,PHW-SA,was characterized by molecular cytological observation and tested for stripe rust resistance in order to evaluate the potential use of the P.huashanica for wheat improvement.The PHW-SA plants were deeply covered with floss in basal spikelet and purple internode,which is characteristic of the P.huashanica parent.Feulgen staining of the somatic metaphases revealed that the chromosome number varied from 51 to 56.At metaphaseⅠ,PHW-SA(2n=56) regularly showed averagely 1.15 univalents and 27.34(24-28) bivalents per cell,and complete chromosome pairing was found in 49.79%of the pollen mother cells(PMCs),indicating a degree of cytological stability.Giemsa-C banding showed that the P.huashanica chromosomes in PHW-SA produced strong bands only in their terminal regions.Genomic in situ hybridization(GISH) analysis suggested that PHW-SA carried 10 entire P. huashanica chromosomes,36 wheat chromosomes and 10 wheat-P,huashanica translocated chromosomes.Seeds storage proteins separated by acid polyacrylamide gel electrophoresis(A-PAGE) and sodium dodecyl sulphate-polyacrylarnide gel electrophoresis(SDS-PAGE) showed that PHW-SA expressed some of P.huashanica specific gliadin and glutenin bands,and some new bands appeared.A survey of diseases resistance revealed that the stripe rust resistance from P.huashanica was totally expressed and powdery mildew was suppressed in the PHW-SA.Random amplified polymorphic DNA(RAPD) analysis verified the presence of DNA of the parents in PHW-SA.The present study showed that the amphiploid PHW-SA could serve as novel sources for transfer of valuable genes to wheat.
引文
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