摘要
油菜属于十字花科芸薹属植物,是世界上四大油料作物之一,也是重要的食用油源和蛋白质饲料来源及重要的工业原料。目前,在我国甘蓝型油菜已经代替了白菜型油菜和芥菜型油菜成为我国主要栽培油菜类型。但是,由于甘蓝型油菜在我国种植时间较晚,遗传基础薄弱,严重制约了我国甘蓝型油菜的育种工作。然而,芸薹属内丰富的遗传资源为甘蓝型油菜种质资源创新提供了条件。通过芸薹属内种间杂交拓宽甘蓝型油菜种质资源,发掘出油菜近缘种中关于产量和农艺性状的优良基因,已经成为改良现有甘蓝型油菜、提高产量的重要手段,也是遗传育种科研工作者研究的主要方向之一。本文利用芥菜型油菜和甘蓝进行种间杂交,采用幼胚培养技术创建异源三倍体,并对其后代进行细胞学、形态学及分子生物学鉴定；对甘蓝型油菜新种质(A+A+C+C+,由芥菜型油菜AjAjBjBj与甘蓝C°C°染色体组部分交换后形成)中半矮秆衍生系10D130进行了株高、角果长度等性状进行了遗传分析。研究结果如下：
1.通过15份芥菜型油菜和6份甘蓝进行种间杂交,并统计30d后残留子房和获得幼胚数,结果表明不同的杂交组合子房残留情况和每100蕾获得幼胚数存在较大差异；对正反交获得幼胚数进行比较,结果表明以芥菜型油菜作母本有利于提高获得幼胚的概率。
2.对184个种间杂种无性系进行形态学鉴定,确认真杂种43份。真杂种表现出较强的生长优势。根据花器官形态和花粉育性将三倍体真杂种分为高度不育(无粉,花粉可染率为0%)、少粉(花粉可染率为5.35%-12.00%)和微粉(花粉可染率为25.3%-47.87%)三种类型。收获时统计不同种间杂种材料自交结实情况,发现种子多表现为干瘪或者半饱满,结实率介于0-3.65%。另外,获得一株染色体加倍植株,该植株花药饱满,每角果结实2-4粒种子。
3.选择其中20份真杂种无性系进行染色体数目调查,发现8份染色体数目为27条,9份染色体数目在27-34之间,3份染色体数目因在不同细胞间变化较大而无法确定,表明杂种细胞在有丝分裂过程中存在非均等分裂,一些细胞存在着部分染色体加倍现象。对细胞减数分裂过程进行观察,结果表明多数花粉母细胞前期联会异常,大多形成单价体和部分二价体；在减数分裂后期I和后期Ⅱ染色体落后严重,多为偏分离。染色体加倍后的植株,部分细胞仍然异常,后期Ⅰ、后期Ⅱ发现有染色体落后现象,但落后染色体数目和发生频率较异源三倍体少。
4.利用芥菜型油菜04K220和羽衣甘蓝04K04种间杂种后代进行系统选育甘蓝型油菜新种质(A+A+C+C+)。研究发现种间杂种(So)通过幼胚挽救获得自交结实无性系(S1),其后代结实率逐代提高。通过2005年到2012年连续选育,共获得122个外部形态类似于甘蓝型油菜的后代衍生系(A+A+C+C+)。选取其中78份后代衍生系(甘蓝型油菜新种质)和2个亲本,利用37对多态性较好的SSR引物进行PCR扩增,结果表明,37对引物共扩出144多态性位点,每对引物平均扩增出3.89条,其中124条为多态性位点,占86.1%；124个位点中,较父本(04K04)和母本(04K22)新增加34个,占总扩增多态性位点的27.42%；丢失9位点,占总扩增多态性位点的7.26%。对2个亲本和78个衍生系进行聚类分析,结果表明种间杂种在多代选育过程中遗传物质发生大量丢失和重排。78个杂种衍生后代品系遗传相似系数在0.65-1.00之间变化,当相似系数在0.72时,可以将所有甘蓝型油菜新种质(A+A+C+C+)分为4个类群。
5.对78份甘蓝型油菜新种质(A+A+C+C+)株高及其相关性状进行调查,结果部分衍生系植株高度低于重庆地区正常油菜植株高度50-60cm,分枝部位低、分枝数目多。对78份新种质的角果长度进行调查,结果该衍生群体内角果长短,部分品系角果长低于4cm。利用近红外对78份新种质的含油量、硫甙和芥酸含量进行测定,其中含油量高于45%的衍生系4个,硫甙低于35umol.g-1的衍生系3个,芥酸含量小于1%的23个,高于55%的2个。
6.10D130是从甘蓝型油菜新种质(A+A+C+C+)中选育出的1个黄籽高油(>45%)半矮杆新种质。其平均株高162cm,比中双11号平均低54cm；有效分枝部位、主花序长、主茎长度和有效分枝间距分别较中双11号减少39.46%、14.8%、22.49%、18.86%；有效分枝数较中双11号增加28.59%。以主基因+多基因混合遗传模型对组合(10D130×中双11)6世代遗传群体(P1、P2、F1、B1、B2和F2)株高及其组成性状进行遗传分析。结果表明,其株高、主茎长、分枝部位、主花序长的遗传均受到1对加性-显性主基因+加性-显性-上位性多基因混合遗传模型控制(D-O模型)。有效分枝间距的最适宜遗传模型为E-1模型(2对加-显-上位性主基因+加-显-上位性多基因混合遗传模型)。该组合有效分枝部位高度、主花序长、有效分枝节间距和有效分枝数与株高均呈显著正相关。从性状表现、遗传来源和影响株高的遗传模型等方面与前人报道的矮秆和半矮秆种质相比,结果显示10D130是一个新型的半矮秆种质资源。
7.10D130和中双11在果身长、角果长和果喙长3个性状差异显著(P<0.05)。采用主+多基因遗传模型对该组合6世代遗传群体(P1、P2、F1、B1、B2和F2)果身长、角果长和果喙长进行遗传分析。结果该杂交组合果身长、角果长和果喙长3个性状均受2对主基因+多基因混合遗传模型控制。其中,果身长最适宜遗传模型为E-0(2对加性-显性-上位性主基因+加性-显性-上位性多基因混合遗传模型)；角果长的最适宜遗传模型为E-1(2对加性-显性-上位性主基因+加性-显性多基因混合遗传模型)；果喙长的最适宜遗传模型为E-3模型(2对加性主基因+加-显多基因混合遗传模型)。此外,果身长、角果长和果喙长3个性状环境因素引起的遗传变异较小,可以在早期世代进行选择。
Rapeseed is one of the major oil crops in the world, which is a kinds of Brassica plant in Cruciferous. It is not only important source of edible oil and protein feed, but also is one of major industrial raw materials. At present, Brassica napus is cultivated widely in China instead of Brassica campestris and Brassica juncea. Thus, weak and narrow genetic foundation of Brassica napus with short history of cultivation in China seriously limited the breeding of Brassica napus. However, on the other hand, abundant genetic resources within Brassica are helpful for creation of germplasm of Brassica napus. Therefore, interspecific hybrids in Brassica are important and efficient method to build and broaden Brassica napus germplasm resources, and discover some helpful genes which can improve quality and yield of rapeseed. This is also one of main research directions of genetics and breeding for rapeseed researcher. Allotriploid were artificially synthesized by culture of immature embryos from interspecific hybridization between Brassica juncea and Brassica oleracea, and the derived lines were identified at cytology, morphology and molecular level. At last, it was used to genetic analysis at the aspect of plant heights and pod lengths for Brassica napus. new germplasm and semi-dwarf10D130(A+A+C+C+), which formed by partly exchange of genome both Brassica juncea (AjAjBjBj) and Brassica oleracea (C°C°). The results are as follows:
1. Interspecific hybridization between15Brassica juncea and6Brassica oleracea were got,30d later, the numbers of residual ovary and immature embryo were recorded, the results showed that there were obviously different number of ovary residual and active immature embryos per100buds in different crosses, and the numbers of immature embryos from positive and negative cross were compared, the results showed that Brassica juncea as female help to improve the probability of immature embryo.
2.43true hybrids were confirmed by identification of and chromosome number among184kinds of hybrid clones. Vegetative of true hybrids showed strong growth advantage, and apparent morphology was more close to the Brassica napus. These triploid hybrids were divided into three types according to floral organ morphology and pollen fertility, including the highly sterile (no powder, pollen contamination rate was0%), less powder (pollen efficiency was5.35%-12.00%) and fine powder (pollen efficiency was25.3%-47.87%), respectively. Seeds from different kinds of hybrids were recorded at harvest periods, the results found most of seeds were empty or semi-full, and seed setting rate change from0to3.65%. In addition, a plant doubled chromosome was got with full anther, and developed2-4seeds per pod.
3. Chromosome numbers among20clones were surveyed, the results found8clones had27chromosomes and9clones had27-34chromosomes, respectively, and others clones could hardly be determined number of chromosome because of great changes among different cell. This indicated that the unequal divisions and chromosome doubling happened during mitosis of some cells. By observing the process of meiosis, the results show that the majority of pollen mother cells early joint abnormalities, mostly formed univalents and some bivalents; due to segregation distortion, chromosomes serious backward during meiosis anaphase Ⅰ and anaphase Ⅱ. For plants of Chromosome doubling, some cells remain chromosome lag during late Ⅰ, anaphase Ⅱ, but the rate and number of lagging chromosomes were less than allotriploid.
4. Brassica napus (A+A+C+C+) were artificially synthesized by systematic selection and breeding of Hybrids between Brassica juncea (04K220) and Brassica napus (04K04). The study showed that the interspecific hybrids (So) selfed to get clones (S1) by rescue of immature embryo, and seed setting rate improved from generation to next generation. Through continuous selection and breeding from2005to2012, there were122descendant lines whose morphologies were similar to be Brassica napus.78of122descendant lines and the two parents conducted PCR amplification by37 polymorphic efficient SSR primers, the results showed that the expansion of144with37pairs of primers, each primer pair amplified by an average of3.89, of which124were polymorphic, accounting for86.1%; comparing with the male parent (04K04) and female parent (04K22),34bands increase and9bands missing among124bands, which accounted for27.42%and7.26%of the amplified polymorphic bands, respectively. cluster analysis among two parents and78lines showed that genetic material from the interspecific hybrid happen great loss and rearrangement during the multi-generation breeding process. Genetic similarity coefficient of78hybrid derivative lines varies between0.65-1.00. When the similarity coefficient is0.72, all hybrid derivative lines and parents can be divided into four groups.
5. Plant heights and pod lengths and their components of78derived lines were surveyed, the results showed that some of derived lines were lower50-60cm plant height, lower mycobacterium site, more branches, shorter pod length than those of normal rapeseed in Chongqing. The oil content, glucosinolate and erucic acid content of78lines were skimmed by using near infrared, in which oil content of4derived lines higher than45%, the glucosinolates content3of them is less than35umol.g, erucic acid content of23lines is less than1%and that2of them lines is higher than55%.
6.10D130is a kind of new and semi-dwarf germplasm with yellow seed coat and high oil contents, which was artificially synthesized to Brassica napus.(A+A+C+C+). average plant height of10D130is162cm in the rapeseed planting areas of Chongqing, which decreased by54cm comparing with conventional elite cultivars of Zhongshuang11. Its branch height, length of main inflorescence, main stem length and valid branch space reduced39.46%,14.8%,22.49%,18.86%comparing with Zhongshuang11, respectively, and numbers of its valid branch rose28.59over Zhongshuang11. These traits above mentioned were significant difference (P<0.05). Six generations (P1, P2, F1, F2, B1, and B2) were produced from a cross between semi-dwarf10D130and Zhongshuang11(ZS11) to analyze inheritances of plant height and related characteristics by the mixed major gene plus polygene inheritance model. The results showed that the plant height, branch height, length of main inflorescence and valid branch space of10D130×ZS11were dominated by a pair of major gene with additive-dominant effects plus polygenes with additive-dominance-epistasis effects (D-0model). The first valid branch space of10D130×ZS11was dominated by two pairs of major gene with additive-dominant-epistasis effects plus polygenes with additive-dominance-epistasis (E-1model). The branch height, length of main inflorescence, valid branch space and number of valid branch were significantly and positively correlated with plant height. Therefore, judged from traits, genetic sources and genetic models,10D130is a different new germplasm with semi-dwarf comparing with previous research.
7. Genetic models of silique body length (SBL), valid silique length (SL) and beak length (BL) were studied in the cross of Zhongshuangll(P1)×10D130(P2), in which difference between P1with long-silique variety and P2with short-silique line is significant (P<0.05). SBL, SL and BL of six populations (P1, P2, F1, F2, B1and B2) were analyzed by joint segregation analysis. The frequency distributions of the F2, BCP1and B2generation exhibited to have multi-modal patterns, indicating the influence of major genes mixed with polygenes. To test and disclose the genetic bases of SBL, SL and BL, the most suitable genetic models were selected among twenty-three genetic models established about three silique traits. The results showed that SBL was well described by E-0model, a case of two additive-dominant-epistasis major gene as well as additive-dominant-epistasis polygenes,. and SL was controlled by two additive-dominant-epistasis major gene as well as additive-dominant polygene (E-1model). In addition, BL was controlled by two additive major genes as well as additive-dominance polygene (E-3). In addition, it is a few little genetic variations of silique body length (SBL), valid silique length (SL) and beak length (BL) caused by environmental factors so as to select and breed in the early generations.
引文
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