小麦phKL和节节麦Ppd-D1基因的遗传评价
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摘要
目前应用于普通小麦(Triticum aestivum L.,2n=6x=42,AABBDD)外源遗传物质转移中的Ph基因突变体ph1b,ph2a和ph2b都是通过人工诱变的方法获得的,这些人工突变体在小麦外源遗传物质转移中已有许多研究。不同于上述人工突变体,四川小麦地方品种开县罗汉麦天然存在phKL基因,其诱导部分同源染色体配对能力类似于Ph2基因突变体,该基因可能与Ph1和Ph2基因突变体有不同的遗传作用机制,可能是一种新型的染色体操作工具材料;光周期不敏感普通小麦在小麦栽培品种中占主导地位,对全世界小麦生产有重要贡献。其中,2D染色体上的光周期基因Ppd-D1对小麦光周期不敏感,发挥了重要作用。在欧洲,多数对光周期不敏感的小麦光周期基因Ppd-D1来源于日本品种Akakomugi。由于仅有极少数的节节麦(Aegilops tauschii Coss.,2n=2x=14,DD)参与了六倍体小麦的起源与进化,节节麦2D染色体上可能存在一些目前普通小麦遗传群体不具有的光周期基因位点。
本文围绕开县罗汉麦天然的phKL基因和节节麦的光周期基因Ppd-D1进行相关研究。利用具有phKL的开县罗汉麦与外源物种Aegilops peregrina(2n=4x=28,UUS~sS~s)远缘杂交,选育稳定株系,并对稳定株系进行分子细胞学鉴定,以评价phKL基因在远缘杂交转移外源基因的实际应用效果;对通过phKL基因导入Ae.peregrina遗传物质的小麦品系进行条锈病抗性鉴定分析,获得新的抗病材料;研究节节麦的光周期基因的遗传多样性,并转入六倍体遗传背景中,评价其应用价值。主要研究结果如下:
1.用含有phKL基因的开县罗汉麦与Ae.peregrina居群13E远缘杂交,杂种用小麦品系3854回交2次后,连续自交选择,获得农艺性状稳定、结实正常、染色体数2n=42的6个品系3121,3131,3141,3161,3171,3181。这6个品系在F3时来自同一个单株。其中,3161,3171,3181在F4时是同一个单株,在F5后分别为3个农艺性状相似的姊妹系。这3个品系与普通小麦类似,软壳、易脱粒,方型穗,植株偏高。其中,3171在成株期抗条锈,而3161和3181感条锈。3121,3131,3141在F4时也是同一个单株,在F5后分别为3个单株的姊妹系。这三个姊妹系的颖壳硬且紧包种子,在室内考种时不易脱粒,只能用手才能剥离,这种颖壳紧包种子而不易脱粒是Ae.peregrina的野生性状。同时,这3个品系与Ae.peregrina居群13E相似之处还表现在:穗型类似,株型分散。从形态上看,3121,3131,3141可能转入了2U或2S~L染色体上的硬壳基因。这三个品系其余主要农艺性状也基本相似。3131和3141在田间无法区分,但是与3121能够根据条锈病的抗性区分开。3131和3141在成株期感条锈,而3121抗条锈。3121的抗条锈特征与13E类似,而且其小麦亲本不抗病,表明抗性来自13E。3121具有的抗病基因与13E的第2同源群硬壳位点无直接关系,可能受不同染色体(或片段)控制。对抗条锈的新品系3121花粉母细胞观察表明,染色体减数分裂过程正常,表明3121是个细胞学上稳定的材料。它与开县罗汉麦杂种F1的绝大多数花粉母细胞配成21个二价体。根据细胞学和农艺性状资料,3121可能是一个具有两个以上易位的易位系。本研究表明,phKL基因在外源基因转移方面有较大的应用价值。但是,通过染色体C带和基因组原位杂交,未成功检测出易位染色体。下一步,需要用更灵敏的分子生物学技术检测出上述品系的易位片段。
2.以开县罗汉麦为母本,13E为父本杂交,将杂种F1中将自交结实的种子,分单株种植,而后选结实率较高单株自交至F9,选出3271,3281,3291,3301,3311等比较稳定的品系。尽管这些品系的穗子比开县罗汉麦偏长,小穗数增多,但是与亲本比较,普遍存在结实率偏低问题。细胞学观察表明,这些品系为细胞学稳定的材料。这些品系结实率不高,可能不是由于细胞学上的减数分裂不正常引起的,而可能是由于导入外源染色体片段不能完全补偿缺失的小麦染色体的遗传物质,致使育性降低。进一步,通过对3271与KL的杂种F1染色体观察,发现绝大多数花粉母细胞配成21个二价体,表明品系3271可能是易位系。对这些品系的高分子量谷蛋白亚基和微卫星(SSR)分子标记分析揭示了新的变异,表明在远缘杂交过程中产生了遗传变异。
3.对56个不同来源节节麦的抽穗期遗传评价表明,节节麦抽穗期存在很大差异。以抽穗期相差7天作为一个间隔,56个节节麦材料被划分为4种类型。类型Ⅰ具有相对短的抽穗期(在169到176天之间)。类型Ⅱ抽穗期在177到184天之间。类型Ⅰ和类型Ⅱ所包含的所有节节麦材料均属于节节麦tauschii亚种。类型Ⅲ抽穗期在185到192天之间。类型Ⅳ抽穗期超过193天。抽穗期与其地理分布相关。分布于我国新疆伊犁河流域和黄河流域的所有节节麦都具有早的抽穗期(类型Ⅰ),它们均属于节节麦tauschii亚种。另一方面,所有strangulata亚种的节节麦只分布于外高加索(亚美尼亚和阿塞拜疆)和伊朗的里海,它们具有晚的抽穗期。三个人工合成六倍体小麦SHW-L1,Syn-SAU-13和Syn-SAU-14的抽穗期分别为182天,189天和189天。它们的共同四倍体供体小麦圆锥麦AS2255的抽穗期为171天,而其三个不同节节麦ssp.tauschii AS60,ssp.tauschii AS2395和ssp.strangulata AS2393的抽穗期分别为173天,195天和195天。由于SHW-L1的抽穗期比Syn-SAU-13和Syn-SAU-14要早7天,推测人工合成六倍体小麦的抽穗期早晚与供体节节麦的抽穗期早晚相关。进一步的遗传分析表明,人工合成六倍体小麦SHW-L1的2D染色体,在SHW-L1和中国春杂种F2的遗传背景下,能缩短抽穗期5天以上。人工合成小麦SHW-L1的2D染色体是由节节麦AS60提供。因此推测,节节麦含有的光周期位点与中国春的ppd-D1位点有差异。由于进化的瓶颈效应,节节麦tauschii亚种具有的光周期位点,可能没有渗入到六倍体小麦中。因此节节麦tauschii亚种AS60中的光周期位点可作为六倍体小麦光周期育种的新资源。
4.前人通过对光周期不敏感和敏感小麦的比较研究发现,光周期不敏感品种的Ppd-D1基因编码区上游序列出现了2089bp缺失,此缺失可能切除了某些调节基因或改变了转录的起始位点,引起Ppd-D1的异常表达,从而启动光周期途径,并最终导致提前抽穗。为了检测光周期不敏感小麦品种Ppd-D1编码区上游存在的2089bp缺失是否也存在节节麦中,我们运用位于此缺失序列之内的PCR扩增引物对8份节节麦扩增,并对产物进行克隆测序。序列比较发现:(1)所有的8份节节麦与光周期敏感普通小麦一样,没有发生2098bp缺失。节节麦未发现该2089bp片段缺失表明,该片段缺失可能是在普通小麦起源之后产生的。由于这8份节节麦的抽穗期存在很大差异,表明节节麦抽穗期存在的差异与这整个2098 bp缺失无直接关系;(2)节节麦的这段序列存在两个不同位置的缺失/插入短序列,分别包含24个碱基(8个氨基酸)和15个碱基(5个氨基酸)。24碱基缺失/插入和临近序列具有MITE(miniature terminal inverted repeat element)转座子特征,该MITE有14 bp不完全的倒位重复序列(CCCATTGGGTATA与GATACCCGATGGG),同时产生了“TA”TSD(target site duplication)重复位点,表明可能是一个MITE活动的结果;(3)存在这两个短序列的节节麦都属于ssp.tauschii,而缺失这两个短序列的节节麦都属于ssp.strangulata。同时,普通小麦也缺失这两个短序列。因此,本研究证明节节麦strangulata亚种是原始普通小麦光周期基因的供体。这也为节节麦strangulata亚种是普通小麦D基因组的供体物种提供了新的证据。(4)根据抽穗期资料,这两个缺失/插入短序列与节节麦的抽穗期早晚并无必然的关系。
5.前人研究表明,普通小麦光周期基因中,Ppd-D1与Ppd-B1,Ppd-A1相比较有一个很大的不同,就是Ppd-D1在其第8外显子中少了16bp的片段。为了研究该16bp片段缺失是否来自于节节麦,我们对11份节节麦材料Ppd-D1基因的第8外显子部分序列克隆测序。序列比较结果表明,与普通小麦不一样,所有的节节麦都没有该16bp片段缺失,这表明该缺失也可能是在普通小麦起源之后产生的。
6.节节麦AS60与普通小麦中国春的光周期基因在控制小麦抽穗早晚方面存在差异,节节麦AS60具有Ppd-D~t1基因的表现型,而中国春具有ppa-D1基因的表现型。但是,AS60并不存在光周期不敏感普通小麦Ppd-D1基因编码区上游序列出现的2089bp缺失。同时,节节麦AS60的Ppd-D~t1基因编码区3'端的1572bp序列与普通小麦中国春相应序列相比,除了上述提到的中国春存在16bp片段缺失之外,仅存在3个SNP位点,表明该段序列差异较小。只有将节节麦AS60的整个Ppd-D~t1基因序列测出来,并与普通小麦比较,才能揭示节节麦AS60与普通小麦中国春的光周期基因在控制小麦抽穗早晚存在差异的遗传原因。
The artificial mutants ph1b, ph2a, and ph2b in common wheat (Triticum aestivum L., 2n=6x=42, AABBDD) have been widely used in alien genetic introgression into wheat. Different from these artificial mutants, Sichuan common wheat landrace Kaixianluohanmai naturally has a recessive gene phKL, which exerts similar strength of mutant in Ph2 locus in wheat-alien hybrids. Gene phKL probably has a different genetic mechanism from mutants in Ph1 and Ph2 loci on controlling homoeologous pairing. Gene phKL may be a new tool for chromosome manipulation; Photoperiod insensitivity is widespread in the world's wheat varieties and acts on a very important role on wheat product. The major source of photoperiod insensitivity in wheat is the Ppd-D1 allele on the short arm of chromosome 2D in common wheat. In the case of European varieties, most hexaploid wheat cultivars insensitive to photoperiod carry a Ppd-D1 allele are derived from the Japanese variety Akakomugi. Because of the evolution bottleneck, much of the genetic variation on Ppd-D1 alleles in the ancestral species Ae. tauschii Coss. (2n=2x=14, DD) may be not represented in common wheat.
In order to evaluate the value of gene phKL on alien genetic introgression, we developed new common wheat lines derived from the hybrids of common wheat landrace Kaixianluohanmai with Aegilops peregrina(2n=4x=28, UUS~SS~S). The new lines were characterized by morphological, cytological, and molecular levels. New common wheat lines with resistance to stripe rust were obtained. Meanwhile, the genetic diversity of Ppd-D1 alleles from Ae. tauschii was evaluated in Ae. tauschii itself or its synthetic hexaploid wheats. The results were as follows:
The hybrids of common wheat landrace Kaixianluohanmai with Aegilops peregrina accession 13E were backcrossed to common wheat elite line 3854 twice. The backcross progenies were repeatly self-crossed and six wheat lines, 3121, 3131, 3141, 3161, 3171, and 3181, were selected. They showed good seed-setting and had 2n=42 chromosomes. The six lines can be traced to a single F3 plant. Of which, the three sister lines 3161,3171, and 3181 were derived from a single F4 plant. They had soft glumes to be easily threshed, square spikes and high plants. Although 3161 and 3181 were susceptible, 3171 was resistant to stripe rust at adult stage. The three sister lines 3121, 3131, and 3141 had similar agronomic and come from a single F4 plant. The three sister lines had tough glumes and were difficult to be threshed, which was inherited from Ae. peregrina. Additionally, they showed speared spike and loose plant-type similar to those of Ae. peregrina. Based on morphological characteristics, the gene for tough glume (Tg) in Ae. peregrina was transferred into wheat lines 3121, 3131, and 3141. Line 3131 and 3141 can not be differentiated each other according to morphological characteristics. However, the two lines were different from line 3121 on resistance to strip rust. Lines 3131 and 3141 were susceptible, while 3121 was resistant to stripe rust at adult stage with a similar resistant characters to Ae. peregrina 13E. Meanwhile, common wheat parents of 3121 was also susceptible to stripe rust at adult stage. Therefore, the resistance should come from Ae. peregrina 13E. The two genes, for resistance to stipe rust and tough glume, may be located on different chromosomes or regions. Pollen-mother-cell (PMC) observation on line 3121 resistant to stripe rust indicated that it was stable in cytology with regular chromosome pairing. The meiotic chromosomes in most PMCs of the 3121×Kaixianluohanmai hybrids showed 21 bivalents. Based on cytological and morphological characteristics, it was suggested that 3121 had more than two translocations. Present study suggested that phKL is useful in alien introgression into wheat. However, further works are still needed to detect Ae. peregrina chromosome fragments existed in above new lines.
From the F9 of hybrids bewteen Kaixianluohanmai and Ae. peregrina 13E, new lines 3271, 3281, 3291, 3301, and 3311were selected. They had more spikelets and longer spike. However, they showed a reduced seed-set rate in comparison with that of Kaixianluohanmai. Because of their regular meiotic process for 42 chromosomes, the lower seed-set may be related to the fail compensation of alien chromosome (fragment) for the loss chromosome (fragment) of wheat. The meiotic chromosomes in most PMCs of the 3171×Kaixianluohanmai hybrids showed 21 bivalents, which suggested that 3121 was a translocation line. Analysis on high-molecular-weight glutenin subunits and microsatellite (SSR) revealed new variations in these new line, which may be induced during the wheat- Ae. peregrina wide cross.
The present study revealed a wide variation for heading time among 56 Ae. tauschii accessions. On the basis of the heading date with an interval for 7 days (a week), the 56 Ae. tauschii accessions were grouped into four types. Types I with relatively short heading date between 169 and 176 days and II with heading date between 177 and 184 days belonged to Ae. tauschii ssp. tauschii. Type III showed heading date between 185 and 192 days. Type IV showed long heading date more than 193 days. The heading date was related to distribution. All the accessions from Yili River valley of Xinjiang, and the middle reaches of the Yellow River in China showed short heading dates (Type I), whereas all the ssp. strangulata accessions were only distributed in Transcaucasia (Armenia, Azerbaijan) and SE Caspian sea in Iran, and they showed very long heading date. The heading date for the same female parent AS2255 of the three synthetic wheats SHW-L1, Syn-SAU-13, and Syn-SAU-14 was 171 days. The heading dates of the three male parents Ae. tauschii ssp. tauschii AS60 for SHW-L1, ssp. tauschii AS2395 for Syn-SAU-13, and ssp. strangulata AS2393 for Syn-SAU-14 were 173, 195, and 195 respectively. The heading dates for synthetic hexaploid wheats SHW-L1, Syn-SAU-13, and Syn-SAU-14 were 182, 189, and 189 days, respectively. These results suggested that heading date of synthetic hexaploid wheats was related to that of Ae. tauschii. Compared to Syn-SAU-13 and Syn-SAU-14, SHW-L1 showed earlier heading time. In contrast, compared to that of AS2255 and AS60, the heading date of SHW-L1 was delayed 11 and 9 days, respectively. However, the heading dates of Syn-SAU-13 and Syn-SAU-14 were shortened in comparison with their corresponding Ae. tauschii parent. Genetic analysis indicated that chromosome 2D of synthetic wheat SHW-L1 showed a major effect on promoting heading time with a reduction of more than 5 days in the hybrid background between SHW-L1 and CS lines. The chromosome 2D of SHW-L1 was provided by Ae. tauschii AS60. Thus, it is postulated that this Ae. tauschii possess a different allele from the ppd-D1 of hexaploid wheat CS. Because of the evolution bottleneck, the allele Ppd-D1 in ssp. tauschii might not be represented in current hexaploid wheat. Therefore, the allele Ppd-D1 in ssp. tauschii AS60 might be used as a new source for hexaploid wheat breeding on photoperiod response.
4. Previous studies indicated that common wheat cultivars with the photoperiod insensitive Ppd-D1 allele had a 2098 bp deletion upstream of the coding region. Cultivars with the photoperiod sensitive ppd-D1 allele did not have the 2098 bp deletion. This deletion was associated with misexpression of the 2D Ppd-D1 gene and expression of the key floral regulator FT (FLOWERING LOCUS T) in short day, showing that photoperiod insensitivity is due to activation of a known photoperiod pathway irrespective of day length. In order to study wehther or not the 2098 bp deletion also exists in Ae. tauschii, primer within the 2098 bp deletion was used in PCR reaction for eight Ae. tauschii accessions. The PCR products were then cloned and sequenced. Sequence comparison indicated that: (1) all the eight Ae. tauschii accessions did not have the 2098 bp deletion as photoperiod sensitive ppd-D1 allele did. The 2098 bp deletion did not appear in Ae. tauschii, indicating that the 2098 bp deletion in the photoperiod insensitive Ppd-D1 allele was produced after the origin of common wheat. The sharp differences on heading time among the eight Ae. tauschii accessions suggested that heading time was not related to the 2098 bp deletion; (2) There were two short sequence insertions/deletions, having 24 bp (8 amino acids) and 15 bp (5 amino acids), within this region. The 24 bp insertions/deletions and adjacent regions seems a result of MITE (miniature terminal inverted repeat element) activation. The MITE insertion had the imperfect 14 bp terminal inverted repeat (TIR, CCCAT(-|T)GGGTATA and GATACCC(-|G)ATGGG), and the MITE insertion produced "TA" target site duplication (TSD); (3) All the Ae. tauschii ssp. tauschii accessions had the two short sequence insertions, wihle all the Ae. tauschii ssp. strangulata accessions and common wheat did not have two short sequence insertions. These result proved that Ae. tauschii ssp. strangulata was the donor of ppd-D1 gene in original common wheat. This also is a new evidence that Ae. tauschii ssp. strangulata was the D-genome donor of common wheat; (4) the two short sequence insertions were not related to heading time of Ae. tauschii.
Previous studies indicated that when compared to the photoperiod gene Ppd-B1 on 2B and Ppd-A1 on 2A, Ppd-D1 on 2D of all common wheat cultivarss that were sequenced had a 16 bp deletion in exon 8. In order to study wehther or not the 16 bp deletion also exists in 2D of Ae. tauschii, primers out of the 16 bp deletion were used in PCR reaction for 11 Ae. tauschii accessions. The PCR products were then cloned and sequenced. Sequence comparison indicated that all the Ae. tauschii accessions did not have the 16 bp deletion, which was different from that of common wheat. These results suggested that the 16 bp deletion in common wheat was also produced after the origin of common wheat.
It was suggested that Ae. tauschii AS60 possess a different allele from the ppd-D1 of hexaploid wheat Chinese Spring. Ae. tauschii AS60 showed the phenotype of Ppd-D~t1, while Chinese Spring showed the phenotype of ppd-D1. However, Ae. tauschii AS60 did not have the 2089 bp deletion as other common wheat with photoperiod insensitive Ppd-D1 allele. Furthermore, compared to that of Chinese Spring, the 1572 bp sequences downstream of Ppd-D~t1 in Ae. tauschii AS60 was conserved. Besides the 16bp deletion in Chinese Spring as above mentioned, there were only three SNPs. In order to further reveal the genetic differences on heading time between Ae. tauschii AS60 and Chinese Spring, the full coding sequence of Ppd-D~t1 in Ae. tauschii AS60 is still needed.
本文围绕开县罗汉麦天然的phKL基因和节节麦的光周期基因Ppd-D1进行相关研究。利用具有phKL的开县罗汉麦与外源物种Aegilops peregrina(2n=4x=28,UUS~sS~s)远缘杂交,选育稳定株系,并对稳定株系进行分子细胞学鉴定,以评价phKL基因在远缘杂交转移外源基因的实际应用效果;对通过phKL基因导入Ae.peregrina遗传物质的小麦品系进行条锈病抗性鉴定分析,获得新的抗病材料;研究节节麦的光周期基因的遗传多样性,并转入六倍体遗传背景中,评价其应用价值。主要研究结果如下:
1.用含有phKL基因的开县罗汉麦与Ae.peregrina居群13E远缘杂交,杂种用小麦品系3854回交2次后,连续自交选择,获得农艺性状稳定、结实正常、染色体数2n=42的6个品系3121,3131,3141,3161,3171,3181。这6个品系在F3时来自同一个单株。其中,3161,3171,3181在F4时是同一个单株,在F5后分别为3个农艺性状相似的姊妹系。这3个品系与普通小麦类似,软壳、易脱粒,方型穗,植株偏高。其中,3171在成株期抗条锈,而3161和3181感条锈。3121,3131,3141在F4时也是同一个单株,在F5后分别为3个单株的姊妹系。这三个姊妹系的颖壳硬且紧包种子,在室内考种时不易脱粒,只能用手才能剥离,这种颖壳紧包种子而不易脱粒是Ae.peregrina的野生性状。同时,这3个品系与Ae.peregrina居群13E相似之处还表现在:穗型类似,株型分散。从形态上看,3121,3131,3141可能转入了2U或2S~L染色体上的硬壳基因。这三个品系其余主要农艺性状也基本相似。3131和3141在田间无法区分,但是与3121能够根据条锈病的抗性区分开。3131和3141在成株期感条锈,而3121抗条锈。3121的抗条锈特征与13E类似,而且其小麦亲本不抗病,表明抗性来自13E。3121具有的抗病基因与13E的第2同源群硬壳位点无直接关系,可能受不同染色体(或片段)控制。对抗条锈的新品系3121花粉母细胞观察表明,染色体减数分裂过程正常,表明3121是个细胞学上稳定的材料。它与开县罗汉麦杂种F1的绝大多数花粉母细胞配成21个二价体。根据细胞学和农艺性状资料,3121可能是一个具有两个以上易位的易位系。本研究表明,phKL基因在外源基因转移方面有较大的应用价值。但是,通过染色体C带和基因组原位杂交,未成功检测出易位染色体。下一步,需要用更灵敏的分子生物学技术检测出上述品系的易位片段。
2.以开县罗汉麦为母本,13E为父本杂交,将杂种F1中将自交结实的种子,分单株种植,而后选结实率较高单株自交至F9,选出3271,3281,3291,3301,3311等比较稳定的品系。尽管这些品系的穗子比开县罗汉麦偏长,小穗数增多,但是与亲本比较,普遍存在结实率偏低问题。细胞学观察表明,这些品系为细胞学稳定的材料。这些品系结实率不高,可能不是由于细胞学上的减数分裂不正常引起的,而可能是由于导入外源染色体片段不能完全补偿缺失的小麦染色体的遗传物质,致使育性降低。进一步,通过对3271与KL的杂种F1染色体观察,发现绝大多数花粉母细胞配成21个二价体,表明品系3271可能是易位系。对这些品系的高分子量谷蛋白亚基和微卫星(SSR)分子标记分析揭示了新的变异,表明在远缘杂交过程中产生了遗传变异。
3.对56个不同来源节节麦的抽穗期遗传评价表明,节节麦抽穗期存在很大差异。以抽穗期相差7天作为一个间隔,56个节节麦材料被划分为4种类型。类型Ⅰ具有相对短的抽穗期(在169到176天之间)。类型Ⅱ抽穗期在177到184天之间。类型Ⅰ和类型Ⅱ所包含的所有节节麦材料均属于节节麦tauschii亚种。类型Ⅲ抽穗期在185到192天之间。类型Ⅳ抽穗期超过193天。抽穗期与其地理分布相关。分布于我国新疆伊犁河流域和黄河流域的所有节节麦都具有早的抽穗期(类型Ⅰ),它们均属于节节麦tauschii亚种。另一方面,所有strangulata亚种的节节麦只分布于外高加索(亚美尼亚和阿塞拜疆)和伊朗的里海,它们具有晚的抽穗期。三个人工合成六倍体小麦SHW-L1,Syn-SAU-13和Syn-SAU-14的抽穗期分别为182天,189天和189天。它们的共同四倍体供体小麦圆锥麦AS2255的抽穗期为171天,而其三个不同节节麦ssp.tauschii AS60,ssp.tauschii AS2395和ssp.strangulata AS2393的抽穗期分别为173天,195天和195天。由于SHW-L1的抽穗期比Syn-SAU-13和Syn-SAU-14要早7天,推测人工合成六倍体小麦的抽穗期早晚与供体节节麦的抽穗期早晚相关。进一步的遗传分析表明,人工合成六倍体小麦SHW-L1的2D染色体,在SHW-L1和中国春杂种F2的遗传背景下,能缩短抽穗期5天以上。人工合成小麦SHW-L1的2D染色体是由节节麦AS60提供。因此推测,节节麦含有的光周期位点与中国春的ppd-D1位点有差异。由于进化的瓶颈效应,节节麦tauschii亚种具有的光周期位点,可能没有渗入到六倍体小麦中。因此节节麦tauschii亚种AS60中的光周期位点可作为六倍体小麦光周期育种的新资源。
4.前人通过对光周期不敏感和敏感小麦的比较研究发现,光周期不敏感品种的Ppd-D1基因编码区上游序列出现了2089bp缺失,此缺失可能切除了某些调节基因或改变了转录的起始位点,引起Ppd-D1的异常表达,从而启动光周期途径,并最终导致提前抽穗。为了检测光周期不敏感小麦品种Ppd-D1编码区上游存在的2089bp缺失是否也存在节节麦中,我们运用位于此缺失序列之内的PCR扩增引物对8份节节麦扩增,并对产物进行克隆测序。序列比较发现:(1)所有的8份节节麦与光周期敏感普通小麦一样,没有发生2098bp缺失。节节麦未发现该2089bp片段缺失表明,该片段缺失可能是在普通小麦起源之后产生的。由于这8份节节麦的抽穗期存在很大差异,表明节节麦抽穗期存在的差异与这整个2098 bp缺失无直接关系;(2)节节麦的这段序列存在两个不同位置的缺失/插入短序列,分别包含24个碱基(8个氨基酸)和15个碱基(5个氨基酸)。24碱基缺失/插入和临近序列具有MITE(miniature terminal inverted repeat element)转座子特征,该MITE有14 bp不完全的倒位重复序列(CCCATTGGGTATA与GATACCCGATGGG),同时产生了“TA”TSD(target site duplication)重复位点,表明可能是一个MITE活动的结果;(3)存在这两个短序列的节节麦都属于ssp.tauschii,而缺失这两个短序列的节节麦都属于ssp.strangulata。同时,普通小麦也缺失这两个短序列。因此,本研究证明节节麦strangulata亚种是原始普通小麦光周期基因的供体。这也为节节麦strangulata亚种是普通小麦D基因组的供体物种提供了新的证据。(4)根据抽穗期资料,这两个缺失/插入短序列与节节麦的抽穗期早晚并无必然的关系。
5.前人研究表明,普通小麦光周期基因中,Ppd-D1与Ppd-B1,Ppd-A1相比较有一个很大的不同,就是Ppd-D1在其第8外显子中少了16bp的片段。为了研究该16bp片段缺失是否来自于节节麦,我们对11份节节麦材料Ppd-D1基因的第8外显子部分序列克隆测序。序列比较结果表明,与普通小麦不一样,所有的节节麦都没有该16bp片段缺失,这表明该缺失也可能是在普通小麦起源之后产生的。
6.节节麦AS60与普通小麦中国春的光周期基因在控制小麦抽穗早晚方面存在差异,节节麦AS60具有Ppd-D~t1基因的表现型,而中国春具有ppa-D1基因的表现型。但是,AS60并不存在光周期不敏感普通小麦Ppd-D1基因编码区上游序列出现的2089bp缺失。同时,节节麦AS60的Ppd-D~t1基因编码区3'端的1572bp序列与普通小麦中国春相应序列相比,除了上述提到的中国春存在16bp片段缺失之外,仅存在3个SNP位点,表明该段序列差异较小。只有将节节麦AS60的整个Ppd-D~t1基因序列测出来,并与普通小麦比较,才能揭示节节麦AS60与普通小麦中国春的光周期基因在控制小麦抽穗早晚存在差异的遗传原因。
The artificial mutants ph1b, ph2a, and ph2b in common wheat (Triticum aestivum L., 2n=6x=42, AABBDD) have been widely used in alien genetic introgression into wheat. Different from these artificial mutants, Sichuan common wheat landrace Kaixianluohanmai naturally has a recessive gene phKL, which exerts similar strength of mutant in Ph2 locus in wheat-alien hybrids. Gene phKL probably has a different genetic mechanism from mutants in Ph1 and Ph2 loci on controlling homoeologous pairing. Gene phKL may be a new tool for chromosome manipulation; Photoperiod insensitivity is widespread in the world's wheat varieties and acts on a very important role on wheat product. The major source of photoperiod insensitivity in wheat is the Ppd-D1 allele on the short arm of chromosome 2D in common wheat. In the case of European varieties, most hexaploid wheat cultivars insensitive to photoperiod carry a Ppd-D1 allele are derived from the Japanese variety Akakomugi. Because of the evolution bottleneck, much of the genetic variation on Ppd-D1 alleles in the ancestral species Ae. tauschii Coss. (2n=2x=14, DD) may be not represented in common wheat.
In order to evaluate the value of gene phKL on alien genetic introgression, we developed new common wheat lines derived from the hybrids of common wheat landrace Kaixianluohanmai with Aegilops peregrina(2n=4x=28, UUS~SS~S). The new lines were characterized by morphological, cytological, and molecular levels. New common wheat lines with resistance to stripe rust were obtained. Meanwhile, the genetic diversity of Ppd-D1 alleles from Ae. tauschii was evaluated in Ae. tauschii itself or its synthetic hexaploid wheats. The results were as follows:
The hybrids of common wheat landrace Kaixianluohanmai with Aegilops peregrina accession 13E were backcrossed to common wheat elite line 3854 twice. The backcross progenies were repeatly self-crossed and six wheat lines, 3121, 3131, 3141, 3161, 3171, and 3181, were selected. They showed good seed-setting and had 2n=42 chromosomes. The six lines can be traced to a single F3 plant. Of which, the three sister lines 3161,3171, and 3181 were derived from a single F4 plant. They had soft glumes to be easily threshed, square spikes and high plants. Although 3161 and 3181 were susceptible, 3171 was resistant to stripe rust at adult stage. The three sister lines 3121, 3131, and 3141 had similar agronomic and come from a single F4 plant. The three sister lines had tough glumes and were difficult to be threshed, which was inherited from Ae. peregrina. Additionally, they showed speared spike and loose plant-type similar to those of Ae. peregrina. Based on morphological characteristics, the gene for tough glume (Tg) in Ae. peregrina was transferred into wheat lines 3121, 3131, and 3141. Line 3131 and 3141 can not be differentiated each other according to morphological characteristics. However, the two lines were different from line 3121 on resistance to strip rust. Lines 3131 and 3141 were susceptible, while 3121 was resistant to stripe rust at adult stage with a similar resistant characters to Ae. peregrina 13E. Meanwhile, common wheat parents of 3121 was also susceptible to stripe rust at adult stage. Therefore, the resistance should come from Ae. peregrina 13E. The two genes, for resistance to stipe rust and tough glume, may be located on different chromosomes or regions. Pollen-mother-cell (PMC) observation on line 3121 resistant to stripe rust indicated that it was stable in cytology with regular chromosome pairing. The meiotic chromosomes in most PMCs of the 3121×Kaixianluohanmai hybrids showed 21 bivalents. Based on cytological and morphological characteristics, it was suggested that 3121 had more than two translocations. Present study suggested that phKL is useful in alien introgression into wheat. However, further works are still needed to detect Ae. peregrina chromosome fragments existed in above new lines.
From the F9 of hybrids bewteen Kaixianluohanmai and Ae. peregrina 13E, new lines 3271, 3281, 3291, 3301, and 3311were selected. They had more spikelets and longer spike. However, they showed a reduced seed-set rate in comparison with that of Kaixianluohanmai. Because of their regular meiotic process for 42 chromosomes, the lower seed-set may be related to the fail compensation of alien chromosome (fragment) for the loss chromosome (fragment) of wheat. The meiotic chromosomes in most PMCs of the 3171×Kaixianluohanmai hybrids showed 21 bivalents, which suggested that 3121 was a translocation line. Analysis on high-molecular-weight glutenin subunits and microsatellite (SSR) revealed new variations in these new line, which may be induced during the wheat- Ae. peregrina wide cross.
The present study revealed a wide variation for heading time among 56 Ae. tauschii accessions. On the basis of the heading date with an interval for 7 days (a week), the 56 Ae. tauschii accessions were grouped into four types. Types I with relatively short heading date between 169 and 176 days and II with heading date between 177 and 184 days belonged to Ae. tauschii ssp. tauschii. Type III showed heading date between 185 and 192 days. Type IV showed long heading date more than 193 days. The heading date was related to distribution. All the accessions from Yili River valley of Xinjiang, and the middle reaches of the Yellow River in China showed short heading dates (Type I), whereas all the ssp. strangulata accessions were only distributed in Transcaucasia (Armenia, Azerbaijan) and SE Caspian sea in Iran, and they showed very long heading date. The heading date for the same female parent AS2255 of the three synthetic wheats SHW-L1, Syn-SAU-13, and Syn-SAU-14 was 171 days. The heading dates of the three male parents Ae. tauschii ssp. tauschii AS60 for SHW-L1, ssp. tauschii AS2395 for Syn-SAU-13, and ssp. strangulata AS2393 for Syn-SAU-14 were 173, 195, and 195 respectively. The heading dates for synthetic hexaploid wheats SHW-L1, Syn-SAU-13, and Syn-SAU-14 were 182, 189, and 189 days, respectively. These results suggested that heading date of synthetic hexaploid wheats was related to that of Ae. tauschii. Compared to Syn-SAU-13 and Syn-SAU-14, SHW-L1 showed earlier heading time. In contrast, compared to that of AS2255 and AS60, the heading date of SHW-L1 was delayed 11 and 9 days, respectively. However, the heading dates of Syn-SAU-13 and Syn-SAU-14 were shortened in comparison with their corresponding Ae. tauschii parent. Genetic analysis indicated that chromosome 2D of synthetic wheat SHW-L1 showed a major effect on promoting heading time with a reduction of more than 5 days in the hybrid background between SHW-L1 and CS lines. The chromosome 2D of SHW-L1 was provided by Ae. tauschii AS60. Thus, it is postulated that this Ae. tauschii possess a different allele from the ppd-D1 of hexaploid wheat CS. Because of the evolution bottleneck, the allele Ppd-D1 in ssp. tauschii might not be represented in current hexaploid wheat. Therefore, the allele Ppd-D1 in ssp. tauschii AS60 might be used as a new source for hexaploid wheat breeding on photoperiod response.
4. Previous studies indicated that common wheat cultivars with the photoperiod insensitive Ppd-D1 allele had a 2098 bp deletion upstream of the coding region. Cultivars with the photoperiod sensitive ppd-D1 allele did not have the 2098 bp deletion. This deletion was associated with misexpression of the 2D Ppd-D1 gene and expression of the key floral regulator FT (FLOWERING LOCUS T) in short day, showing that photoperiod insensitivity is due to activation of a known photoperiod pathway irrespective of day length. In order to study wehther or not the 2098 bp deletion also exists in Ae. tauschii, primer within the 2098 bp deletion was used in PCR reaction for eight Ae. tauschii accessions. The PCR products were then cloned and sequenced. Sequence comparison indicated that: (1) all the eight Ae. tauschii accessions did not have the 2098 bp deletion as photoperiod sensitive ppd-D1 allele did. The 2098 bp deletion did not appear in Ae. tauschii, indicating that the 2098 bp deletion in the photoperiod insensitive Ppd-D1 allele was produced after the origin of common wheat. The sharp differences on heading time among the eight Ae. tauschii accessions suggested that heading time was not related to the 2098 bp deletion; (2) There were two short sequence insertions/deletions, having 24 bp (8 amino acids) and 15 bp (5 amino acids), within this region. The 24 bp insertions/deletions and adjacent regions seems a result of MITE (miniature terminal inverted repeat element) activation. The MITE insertion had the imperfect 14 bp terminal inverted repeat (TIR, CCCAT(-|T)GGGTATA and GATACCC(-|G)ATGGG), and the MITE insertion produced "TA" target site duplication (TSD); (3) All the Ae. tauschii ssp. tauschii accessions had the two short sequence insertions, wihle all the Ae. tauschii ssp. strangulata accessions and common wheat did not have two short sequence insertions. These result proved that Ae. tauschii ssp. strangulata was the donor of ppd-D1 gene in original common wheat. This also is a new evidence that Ae. tauschii ssp. strangulata was the D-genome donor of common wheat; (4) the two short sequence insertions were not related to heading time of Ae. tauschii.
Previous studies indicated that when compared to the photoperiod gene Ppd-B1 on 2B and Ppd-A1 on 2A, Ppd-D1 on 2D of all common wheat cultivarss that were sequenced had a 16 bp deletion in exon 8. In order to study wehther or not the 16 bp deletion also exists in 2D of Ae. tauschii, primers out of the 16 bp deletion were used in PCR reaction for 11 Ae. tauschii accessions. The PCR products were then cloned and sequenced. Sequence comparison indicated that all the Ae. tauschii accessions did not have the 16 bp deletion, which was different from that of common wheat. These results suggested that the 16 bp deletion in common wheat was also produced after the origin of common wheat.
It was suggested that Ae. tauschii AS60 possess a different allele from the ppd-D1 of hexaploid wheat Chinese Spring. Ae. tauschii AS60 showed the phenotype of Ppd-D~t1, while Chinese Spring showed the phenotype of ppd-D1. However, Ae. tauschii AS60 did not have the 2089 bp deletion as other common wheat with photoperiod insensitive Ppd-D1 allele. Furthermore, compared to that of Chinese Spring, the 1572 bp sequences downstream of Ppd-D~t1 in Ae. tauschii AS60 was conserved. Besides the 16bp deletion in Chinese Spring as above mentioned, there were only three SNPs. In order to further reveal the genetic differences on heading time between Ae. tauschii AS60 and Chinese Spring, the full coding sequence of Ppd-D~t1 in Ae. tauschii AS60 is still needed.
引文
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