甘蓝籽粒色泽的遗传及其与甘蓝型油菜的比较
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摘要
黄色籽粒性状广泛存在于十字花科芸薹属植物中。白菜型油菜、芥菜型油菜,尤其是甘蓝型油菜作为食用植物油的主要来源而被广泛种植,对籽粒色泽早已进行了深入系统的研究。而作为甘蓝型油菜祖先亲本种之一的甘蓝,由于一直以来主要用作食用蔬菜和观赏花卉栽培,研究者对其种子性状并未给予应有的关注。
综合分析国内外的研究现状发现,在油菜及其近缘物种籽粒色泽的遗传研究方面至少存在两个主要问题或不足：(1)籽粒色泽的遗传研究主要集中在白菜型油菜、芥菜型油菜和甘蓝型油菜,而有关埃塞俄比亚芥和甘蓝的研究很少,这使得育种家难于充分、有效地利用C基因组的黄籽基因来选育和改良甘蓝型油菜的籽粒色泽,从而极大地限制了甘蓝型黄籽油菜的育种进程；(2)绝大多数研究对籽粒色泽的观察和统计都采用目测法,并将籽粒色泽看成质量性状,应用孟德尔的分类统计法进行遗传分析,而应用其他度量方法和数量性状遗传分析方法的研究很少；由于目测法带有观察者的主观性和不确定性(有时还存在系统误差),不同的研究者采用的分级标准不同,分类统计的结果也不同,这使得遗传分析结论的准确性和可靠性大大降低。同时,也未见与籽粒色泽直接相关的甘蓝种皮色素和环境条件对甘蓝籽粒色泽影响的研究报道。
重庆市油菜工程技术研究中心于1994年首次在观赏植物羽衣甘蓝中发现黄籽单株,进而选育出遗传稳定的黄籽株系材料。这是国际上报道的首例甘蓝黄籽材料。本论文以黄籽甘蓝品系为核心材料,研究了甘蓝籽粒色泽的遗传特性、主要农艺措施和基因型×环境互作对甘蓝籽粒色泽的影响以及种子发育过程中甘蓝种皮主要色素含量的动态变化、甘蓝成熟种子种皮色素含量的差异,并与甘蓝型油菜进行了对比分析。开展这项研究不仅可以扩展芸薹属植物籽粒色泽遗传研究的领域和广度,而且所获得的研究结果,一方面将大大地加深对黄籽甘蓝型油菜籽粒色泽遗传规律的认识,另一方面将为充分利用甘蓝的C基因组黄籽基因奠定基础,推动黄籽甘蓝型油菜的育种进程。因此,本项研究具有重要的理论意义和实践价值。
1.甘蓝籽粒色泽的遗传分析
以籽粒色泽不同的羽衣甘蓝、结球甘蓝和白花芥蓝为材料配制杂交组合,构建P1、P2、F1、B1、B2和F2等6个世代群体,将籽粒色泽看作数量性状,采用自主开发的数字图像分析法度量甘蓝的籽粒色泽,利用植物数量性状主基因+多基因遗传体系多世代联合分析方法,建立主基因+多基因混合遗传模型,研究甘蓝籽粒色泽的遗传特性,包括黄色与非黄色的显隐性关系、控制基因对数、基因作用方式、母体效应和遗传率等。主要结论如下：
(1)甘蓝籽粒色泽的遗传表现为母体效应,杂交当代的籽粒色泽与杂交母本自交种子相同；F1代籽粒色泽总体表现较深,黄色对非黄色为隐性。
(2)甘蓝籽粒色泽的遗传受1-2对主基因和微效多基因的共同控制,主基因和多基因均具有加性效应和显性效应,或同时具有上位性效应；主基因为部分显性或完全显性,2对主基因的效应相同或不同；多基因为部分显性、完全显性或超显性；3个分离世代的主基因遗传率均大于多基因遗传率,主基因+多基因遗传率较高。甘蓝籽粒色泽的遗传较为复杂,不同的杂交组合具有不同的基因效应,表现为不同的遗传模型。
(3)甘蓝籽粒色泽的表现受环境因素的影响也较大。
2.主要农艺措施和基因型×环境互作对甘蓝籽粒色泽的影响
以籽粒色泽不同的羽衣甘蓝、结球甘蓝和白花芥蓝为材料,采用裂裂区设计和随机区组设计研究施氮量、种植密度、播种期和收获期等主要农艺措施和基因型×环境互作对甘蓝籽粒色泽的影响以及甘蓝籽粒色泽的表现对环境变化的稳定性。主要结论如下：
(1)施氮量、播种期和收获期对甘蓝籽粒色泽的影响因基因型不同而异,对黑籽基因型没有明显的影响,而极显著地影响黄籽基因型的籽粒色泽；黄籽基因型籽粒色泽随施氮量的增加逐渐变浅,提早或推迟播种都会使黄籽基因型籽粒色泽加深,收获过迟会极显著地降低黄籽基因型的黄色程度。种植密度对甘蓝籽粒色泽没有明显的影响。
(2)基因型×环境互作对甘蓝籽粒色泽也有极显著的影响,不同基因型对环境变化的反应不同,表现出稳定性的差异,尤其是黄籽基因型。
3.甘蓝种子发育过程中种皮色素含量的动态变化
以遗传来源不同的3对具有相同遗传背景的黄籽和黑籽羽衣甘蓝为材料研究种子发育过程中甘蓝种皮主要色素(叶绿素、类胡萝卜素、花色素、黑色素和类黄酮)含量的动态变化和甘蓝成熟种子种皮色素含量的差异。主要结论如下：
(1)在甘蓝种子发育过程中,除类胡萝卜素外,种皮叶绿素、花色素、黑色素和类黄酮的含量在不同色泽籽粒种皮中均存在极显著的差异；不同发育时期5种色素的含量都有极显著的变化,叶绿素、花色素和类黄酮的含量均呈现出先升高、后降低的变化趋势,黑籽种皮类胡萝卜素含量单调上升而黄籽先降后升,种皮黑色素含量从开花后40天起迅速增高,黑籽与黄籽间的差异也迅速增大；种子成熟时,黑籽和黄籽种皮类胡萝卜素含量基本相同,而黑籽种皮叶绿素、花色素、黑色素和类黄酮的含量显著或极显著高于遗传背景相同的黄籽；种子发育过程中种皮5种色素含量的变化规律在黄籽和黑籽间具有明显的差异。
(2)甘蓝种子收获脱粒后,种皮叶绿素、花色素和类黄酮的含量还会继续下降,而类胡萝卜素和黑色素的含量继续上升；风干成熟种子黑籽种皮中5种色素含量均明显高于遗传背景相同的黄籽种皮；相对于黄籽而言,黑籽种皮中花色素含量最高,其次是黑色素和叶绿素,类胡萝卜素和类黄酮较小。引起甘蓝风干成熟种子籽粒色泽差异的主要色素是花色素,其次是黑色素和叶绿素,类胡萝卜素和类黄酮也有一定的作用。
4.甘蓝与甘蓝型油菜籽粒色泽遗传的对比分析
通过文献研究法,对比分析甘蓝和甘蓝型油菜在籽粒色泽的遗传特点、主要农艺措施和基因型×环境互作对籽粒色泽的影响、籽粒色泽对环境变化的稳定性、种子发育过程中种皮主要色素的动态变化以及成熟种子种皮色素含量差异等方面的异同。主要结论如下：
(1)甘蓝籽粒色泽的遗传特点与甘蓝型油菜相似,在黄籽性状显隐性、控制基因对数、基因作用方式、母体效应和遗传率等方面都没有明显的差异。
(2)除种植密度外,播种期、收获期和施氮量等主要农艺措施对甘蓝籽粒色泽的影响与甘蓝型油菜基本相同；基因型×环境互作效应对甘蓝籽粒色泽的影响与甘蓝型油菜类似,甘蓝籽粒色泽的稳定性与甘蓝型油菜也相似。
(3)种子发育过程中,除种皮花色素含量最大值出现时间略有不同外,叶绿素、类胡萝卜素、黑色素和类黄酮的变化动态在甘蓝和甘蓝型油菜间没有明显的不同；除甘蓝成熟种子黄、黑籽种皮叶绿素含量之间的差异与甘蓝型油菜不同外,其余4种色素甘蓝与甘蓝型油菜相同；甘蓝成熟种子黄籽与黑籽种皮色素含量差异最大的是花色素,其次是黑色素和叶绿素,而甘蓝型油菜却是黑色素和花色素。
(4)可以认为,甘蓝型油菜黄籽性状的数量性状遗传表现很可能是由来自甘蓝的C基因组上的基因引起的。
Yellow seed is a very common trait of Brassica species in Cruciferae family. Brassica campestris L., Brassica juncea Coss, and Brassica napus L. are widely planted as the main source of edible vegetable oil, and their seed colour has been thoroughly studied. However, Brassica oleracea L., one of the ancestor species of B. napus, was mainly cultivated as edible vegetables and ornamental flowers, and its seed trait has not been payed attention by researchers.
With the current research situations being comprehensively analyzed, at least two main problems or defects were found in seed colour inheritance research of rapeseed and its close species. Firstly, inheritance researches on seed colour mostly focused on B. campestris, B. juncea and B. napus, a few studies were conducted on B. carinata and B. oleraeca. This makes it difficult to effectively use the yellow seed genes of C genome to improve the seed colour of B. napus, and greatly limits the breeding process of yellow-seeded B. napus. Secondly, the seed colour was investigated by using eyes observation and considered as a quality trait and applying the Mendel's classification statistic method in most of heredity researches, whereas other measurement and quantitative trait genetic analysis was seldom used to study seed colour. The observer's subjectivity and uncertainty (and sometimes systematic error) and the different classification standard greatly reduced the accuracy and reliability of genetic analysis results. Meanwhile, none reports were found about seed coat pigment related directly with seed colour and the environment influence on seed colour in B. oleracea.
For the first time, Chongqing Engineering Research Center for Rapeseed found three yellow-seeded plants of ornamental flower B. oleracea var. acephala DC. in1994, and then bred yellow-seeded lines with genetic stability. This was the first report about yellow-seeded B. oleracea in the world. In the present study, these yellow-seeded lines are used as core materials, and the main objectives are to analyze the genetic characteristics of seed colour, the influences of major agronomic measures and genotype×environment interaction on seed colour, the dynamic changes of main pigment contents of seed coat during the process of seed development, and the differences of seed coat pigment contents of B. oleracea mature seed, and to comparatively analyze these respects between B. oleracea and B. napus. This study might enlarge the research field and scope on the inheritance of seed colour in Brassica species. The results will not only greatly deepen our insights into the inheritance law of seed colour in yellow-seeded B. napus, but also lay a foundation for full use of yellow seed genes of C genome from B. oleracea and promote the breeding process of yellow-seeded B. napus. Therefore, this research has important theoretical significance and practical value.
1. Genetic analysis on seed colour in B. oleracea
B. oleracea var. acephala DC., B. oleracea var. capitata L. and B. oleracea var. alboglabra Bailey with different seed colours were used as materials to generate hybrid combinations and the derived populations P1, P2, F1, B1, B2and F2. The seed colour of B. oleracea was considered as quantitative trait and measured by the digital image analysis method which was developed in our labratory. The multiple generation conjoint analysis of major gene plus polygene system of quantitative trait was conducted to establish the mixed genetic model and analyze the hereditary characteristics of seed colour in B. oleracea, including the dominant or recessive gene of yellow seed trait, the number and effect of seed colour gene, as well as maternal effect and heritability. The main conclusions are as followings:
(1) The inheritance of B. oleracea seed colour presented maternal effect and the immediate hybrid seeds had the same colour as the self-pollinated seeds of its female parent. The seed colour of F1generation is darker in general, and yellow seed was recessive trait over the other seed.
(2) The seed colour of B. oleracea was commonly controlled by one or two major gene(s) and minor polygenes. Both the major gene and minor polygenes had additive effect, dominant effect and sometimes had epistatic effect. The major gene was partial dominant or complete dominant and the two pairs of major genes had the same effects in one hybrid combination and had the different effects in other combinations. The polygenes were partial dominant, complete dominant or superdominance in all combinations. Heritabilities of major gene were greater than those of polygenes in the three segregating generations. The estimate values of heritability of major gene and polygenes were relatively high. These results suggested that the inheritance of B. oleracea seed colour was a considerably complex system, and the effects of seed colour genes were quite different in different hybrid combinations and belonged to different genetic models.
(3) The performance of B. oleracea seed colour could be affected by enviromental factors in a certain degree.
2. The effects of agronomic measures and genotype×environment interaction on B. oleracea seed colour
B. oleracea var. acephala, B. oleracea var. capitata and B. oleracea var. alboglabra with different seed colours were used as materials and split-split plot design and randomized block design were adopted to investigate the effects of different agronomic measures, including the amount of nitrogen fertilizer, planting density, sowing and harvest time, and genotype×environment interaction on seed colour, as well as the performance stability of seed colours to the environmental changes in B. oleracea. The main conclusions are as followings:
(1) The effects of the amount of nitrogen fertilizer, sowing and harvest time on seed colour varied with genotypes in B. oleracea. No obvious effects on the seed colour of black-seeded genotypes and very significant influences on the yellow-seeded genotypes were found. The seed colour of yellow-seeded genotypes would gradually become lighter with the amount increase of nitrogen fertilizer. The early or late sowing time would make the seed colour of yellow-seeded genotypes dark. The too late harvest time could extremely significantly reduce the seed yellow degree of yellow-seeded genotypes. Planting density had no apparent effects on B. oleracea seed colour.
(2) The interaction of genotype×environment significantly affected B. oleracea seed colour and different genotypes had different reactions to the environment changes, especially in the yellow-seeded genotypes.
3. The dynamic changes of seed coat pigment content during seed development process in B. oleracea
Three pairs of B. oleracea var. acephala yellow-and black-seeded lines with the same genetic background from different sources were used to analyze the dynamic changes of main seed coat pigment content (chlorophyll, carotenoid, anthocyanidin, melanin and flavonoid) during the seed development process and the differences of seed coat pigment content in mature seed. The main conclusions are as followings:
(1) In the seed development process of B. oleracea, the contents of chlorophyll, anthocyanidin, melanin and flavonoid except carotenoid showed significant differences in seed coat with different colour and all the five pigment contents varied significantly at different development stages. The chlorophyll, anthocyanidin and flavonoid presented a change trend from increasing to decreasing. The carotenoid contents of black seed coat monotonicly increased with the seed development, whereas those of yellow seed coat decreased at the early stage and then increased. Melanin contents increased sharply from40days after flowering and the differences of melanin content enlarged rapidly between black and yellow seed coat. The carotenoid contents of yellow and black seed coat were almost at the same level in mature seed, whereas the contents of chlorophyll, anthocyanidin, melanin and flavonoid of black seed coat were extremely significantly or significantly higher than those of yellow seed coat which had the same genetic background. The different change tendencies of the five pigment contents existed between yellow and black seed coat in the process of seed development.
(2) The contents of chlorophyll, anthocyanidin and flavonoid of seed coat would continue to decline in B. oleracea, whereas the contents of carotenoid and melanin continue to rise after harvest and threshing. The five pigment contents of black seed coat were significantly higher than those of yellow seed coat with the same genetic background in the air-drying mature seed. Comparing with yellow seed coat, the anthocyanidin content was the highest in black seed coat, the contents of melanin and chlorophyll were next, and the contents of carotenoid and flavonoid were the last. Therefore, in air-drying mature seed coat of B. oleracea, the main pigment causing the differences of seed colour was anthocyanidin, next melanin and chlorophyll, and then carotenoid and flavonoid.
4. Comparative analysis of seed colour inheritance in B. oleracea and B. napus
The means of literature research were used to compare and analyze the genetic characteristics of seed colour of B. oleracea and B. napus, the effects of major agronomic measures and genotype×environment interaction on seed colour, the stability of seed colour to the environmental changes, the dynamic changes of main pigment content of seed coat during seed development process and the differences of pigment content in mature seed coat, etc. The main conclusions are as followings:
(1) The hereditary characteristics of B. oleracea seed colour were similar to B. napus and there were no obvious differences in the dominant or recessive gene of yellow seed trait, the number and effect of seed colour gene, as well as maternal effect and heritability, etc.
(2) Except planting density, the effects of major agronomic measures, such as sowing and harvest time and the amount of nitrogen fertilizer, etc, on seed colour were relatively similar between B. oleracea and B. napus. The genotype×environment interaction effect on B. oleracea seed colour and the environmental stability of B. oleracea seed colour were similar to B. napus.
(3) During seed development process, in addition to a slight difference of peak date of anthocyanidin content, the dynamic changes of chlorophyll, carotenoid, melanin and flavonoids were no significant differences between B. oleracea and B. napus seed coat. Between yellow and black mature seed coat, the difference of chlorophyll content in B. oleracea was different with B. napus, the rest four pigments were the same. The largest difference between yellow and black mature seed coat was anthocyanidin content in B. oleracea, followed by melanin and chlorophyll, however they were melanin and anthocyanidin in B. napus.
(4) It suggested that quantitative genetic characteristics of yellow seed trait of B. napus was likely to be resulted from the genes on the C genome come from B. oleracea.
综合分析国内外的研究现状发现,在油菜及其近缘物种籽粒色泽的遗传研究方面至少存在两个主要问题或不足：(1)籽粒色泽的遗传研究主要集中在白菜型油菜、芥菜型油菜和甘蓝型油菜,而有关埃塞俄比亚芥和甘蓝的研究很少,这使得育种家难于充分、有效地利用C基因组的黄籽基因来选育和改良甘蓝型油菜的籽粒色泽,从而极大地限制了甘蓝型黄籽油菜的育种进程；(2)绝大多数研究对籽粒色泽的观察和统计都采用目测法,并将籽粒色泽看成质量性状,应用孟德尔的分类统计法进行遗传分析,而应用其他度量方法和数量性状遗传分析方法的研究很少；由于目测法带有观察者的主观性和不确定性(有时还存在系统误差),不同的研究者采用的分级标准不同,分类统计的结果也不同,这使得遗传分析结论的准确性和可靠性大大降低。同时,也未见与籽粒色泽直接相关的甘蓝种皮色素和环境条件对甘蓝籽粒色泽影响的研究报道。
重庆市油菜工程技术研究中心于1994年首次在观赏植物羽衣甘蓝中发现黄籽单株,进而选育出遗传稳定的黄籽株系材料。这是国际上报道的首例甘蓝黄籽材料。本论文以黄籽甘蓝品系为核心材料,研究了甘蓝籽粒色泽的遗传特性、主要农艺措施和基因型×环境互作对甘蓝籽粒色泽的影响以及种子发育过程中甘蓝种皮主要色素含量的动态变化、甘蓝成熟种子种皮色素含量的差异,并与甘蓝型油菜进行了对比分析。开展这项研究不仅可以扩展芸薹属植物籽粒色泽遗传研究的领域和广度,而且所获得的研究结果,一方面将大大地加深对黄籽甘蓝型油菜籽粒色泽遗传规律的认识,另一方面将为充分利用甘蓝的C基因组黄籽基因奠定基础,推动黄籽甘蓝型油菜的育种进程。因此,本项研究具有重要的理论意义和实践价值。
1.甘蓝籽粒色泽的遗传分析
以籽粒色泽不同的羽衣甘蓝、结球甘蓝和白花芥蓝为材料配制杂交组合,构建P1、P2、F1、B1、B2和F2等6个世代群体,将籽粒色泽看作数量性状,采用自主开发的数字图像分析法度量甘蓝的籽粒色泽,利用植物数量性状主基因+多基因遗传体系多世代联合分析方法,建立主基因+多基因混合遗传模型,研究甘蓝籽粒色泽的遗传特性,包括黄色与非黄色的显隐性关系、控制基因对数、基因作用方式、母体效应和遗传率等。主要结论如下：
(1)甘蓝籽粒色泽的遗传表现为母体效应,杂交当代的籽粒色泽与杂交母本自交种子相同；F1代籽粒色泽总体表现较深,黄色对非黄色为隐性。
(2)甘蓝籽粒色泽的遗传受1-2对主基因和微效多基因的共同控制,主基因和多基因均具有加性效应和显性效应,或同时具有上位性效应；主基因为部分显性或完全显性,2对主基因的效应相同或不同；多基因为部分显性、完全显性或超显性；3个分离世代的主基因遗传率均大于多基因遗传率,主基因+多基因遗传率较高。甘蓝籽粒色泽的遗传较为复杂,不同的杂交组合具有不同的基因效应,表现为不同的遗传模型。
(3)甘蓝籽粒色泽的表现受环境因素的影响也较大。
2.主要农艺措施和基因型×环境互作对甘蓝籽粒色泽的影响
以籽粒色泽不同的羽衣甘蓝、结球甘蓝和白花芥蓝为材料,采用裂裂区设计和随机区组设计研究施氮量、种植密度、播种期和收获期等主要农艺措施和基因型×环境互作对甘蓝籽粒色泽的影响以及甘蓝籽粒色泽的表现对环境变化的稳定性。主要结论如下：
(1)施氮量、播种期和收获期对甘蓝籽粒色泽的影响因基因型不同而异,对黑籽基因型没有明显的影响,而极显著地影响黄籽基因型的籽粒色泽；黄籽基因型籽粒色泽随施氮量的增加逐渐变浅,提早或推迟播种都会使黄籽基因型籽粒色泽加深,收获过迟会极显著地降低黄籽基因型的黄色程度。种植密度对甘蓝籽粒色泽没有明显的影响。
(2)基因型×环境互作对甘蓝籽粒色泽也有极显著的影响,不同基因型对环境变化的反应不同,表现出稳定性的差异,尤其是黄籽基因型。
3.甘蓝种子发育过程中种皮色素含量的动态变化
以遗传来源不同的3对具有相同遗传背景的黄籽和黑籽羽衣甘蓝为材料研究种子发育过程中甘蓝种皮主要色素(叶绿素、类胡萝卜素、花色素、黑色素和类黄酮)含量的动态变化和甘蓝成熟种子种皮色素含量的差异。主要结论如下：
(1)在甘蓝种子发育过程中,除类胡萝卜素外,种皮叶绿素、花色素、黑色素和类黄酮的含量在不同色泽籽粒种皮中均存在极显著的差异；不同发育时期5种色素的含量都有极显著的变化,叶绿素、花色素和类黄酮的含量均呈现出先升高、后降低的变化趋势,黑籽种皮类胡萝卜素含量单调上升而黄籽先降后升,种皮黑色素含量从开花后40天起迅速增高,黑籽与黄籽间的差异也迅速增大；种子成熟时,黑籽和黄籽种皮类胡萝卜素含量基本相同,而黑籽种皮叶绿素、花色素、黑色素和类黄酮的含量显著或极显著高于遗传背景相同的黄籽；种子发育过程中种皮5种色素含量的变化规律在黄籽和黑籽间具有明显的差异。
(2)甘蓝种子收获脱粒后,种皮叶绿素、花色素和类黄酮的含量还会继续下降,而类胡萝卜素和黑色素的含量继续上升；风干成熟种子黑籽种皮中5种色素含量均明显高于遗传背景相同的黄籽种皮；相对于黄籽而言,黑籽种皮中花色素含量最高,其次是黑色素和叶绿素,类胡萝卜素和类黄酮较小。引起甘蓝风干成熟种子籽粒色泽差异的主要色素是花色素,其次是黑色素和叶绿素,类胡萝卜素和类黄酮也有一定的作用。
4.甘蓝与甘蓝型油菜籽粒色泽遗传的对比分析
通过文献研究法,对比分析甘蓝和甘蓝型油菜在籽粒色泽的遗传特点、主要农艺措施和基因型×环境互作对籽粒色泽的影响、籽粒色泽对环境变化的稳定性、种子发育过程中种皮主要色素的动态变化以及成熟种子种皮色素含量差异等方面的异同。主要结论如下：
(1)甘蓝籽粒色泽的遗传特点与甘蓝型油菜相似,在黄籽性状显隐性、控制基因对数、基因作用方式、母体效应和遗传率等方面都没有明显的差异。
(2)除种植密度外,播种期、收获期和施氮量等主要农艺措施对甘蓝籽粒色泽的影响与甘蓝型油菜基本相同；基因型×环境互作效应对甘蓝籽粒色泽的影响与甘蓝型油菜类似,甘蓝籽粒色泽的稳定性与甘蓝型油菜也相似。
(3)种子发育过程中,除种皮花色素含量最大值出现时间略有不同外,叶绿素、类胡萝卜素、黑色素和类黄酮的变化动态在甘蓝和甘蓝型油菜间没有明显的不同；除甘蓝成熟种子黄、黑籽种皮叶绿素含量之间的差异与甘蓝型油菜不同外,其余4种色素甘蓝与甘蓝型油菜相同；甘蓝成熟种子黄籽与黑籽种皮色素含量差异最大的是花色素,其次是黑色素和叶绿素,而甘蓝型油菜却是黑色素和花色素。
(4)可以认为,甘蓝型油菜黄籽性状的数量性状遗传表现很可能是由来自甘蓝的C基因组上的基因引起的。
Yellow seed is a very common trait of Brassica species in Cruciferae family. Brassica campestris L., Brassica juncea Coss, and Brassica napus L. are widely planted as the main source of edible vegetable oil, and their seed colour has been thoroughly studied. However, Brassica oleracea L., one of the ancestor species of B. napus, was mainly cultivated as edible vegetables and ornamental flowers, and its seed trait has not been payed attention by researchers.
With the current research situations being comprehensively analyzed, at least two main problems or defects were found in seed colour inheritance research of rapeseed and its close species. Firstly, inheritance researches on seed colour mostly focused on B. campestris, B. juncea and B. napus, a few studies were conducted on B. carinata and B. oleraeca. This makes it difficult to effectively use the yellow seed genes of C genome to improve the seed colour of B. napus, and greatly limits the breeding process of yellow-seeded B. napus. Secondly, the seed colour was investigated by using eyes observation and considered as a quality trait and applying the Mendel's classification statistic method in most of heredity researches, whereas other measurement and quantitative trait genetic analysis was seldom used to study seed colour. The observer's subjectivity and uncertainty (and sometimes systematic error) and the different classification standard greatly reduced the accuracy and reliability of genetic analysis results. Meanwhile, none reports were found about seed coat pigment related directly with seed colour and the environment influence on seed colour in B. oleracea.
For the first time, Chongqing Engineering Research Center for Rapeseed found three yellow-seeded plants of ornamental flower B. oleracea var. acephala DC. in1994, and then bred yellow-seeded lines with genetic stability. This was the first report about yellow-seeded B. oleracea in the world. In the present study, these yellow-seeded lines are used as core materials, and the main objectives are to analyze the genetic characteristics of seed colour, the influences of major agronomic measures and genotype×environment interaction on seed colour, the dynamic changes of main pigment contents of seed coat during the process of seed development, and the differences of seed coat pigment contents of B. oleracea mature seed, and to comparatively analyze these respects between B. oleracea and B. napus. This study might enlarge the research field and scope on the inheritance of seed colour in Brassica species. The results will not only greatly deepen our insights into the inheritance law of seed colour in yellow-seeded B. napus, but also lay a foundation for full use of yellow seed genes of C genome from B. oleracea and promote the breeding process of yellow-seeded B. napus. Therefore, this research has important theoretical significance and practical value.
1. Genetic analysis on seed colour in B. oleracea
B. oleracea var. acephala DC., B. oleracea var. capitata L. and B. oleracea var. alboglabra Bailey with different seed colours were used as materials to generate hybrid combinations and the derived populations P1, P2, F1, B1, B2and F2. The seed colour of B. oleracea was considered as quantitative trait and measured by the digital image analysis method which was developed in our labratory. The multiple generation conjoint analysis of major gene plus polygene system of quantitative trait was conducted to establish the mixed genetic model and analyze the hereditary characteristics of seed colour in B. oleracea, including the dominant or recessive gene of yellow seed trait, the number and effect of seed colour gene, as well as maternal effect and heritability. The main conclusions are as followings:
(1) The inheritance of B. oleracea seed colour presented maternal effect and the immediate hybrid seeds had the same colour as the self-pollinated seeds of its female parent. The seed colour of F1generation is darker in general, and yellow seed was recessive trait over the other seed.
(2) The seed colour of B. oleracea was commonly controlled by one or two major gene(s) and minor polygenes. Both the major gene and minor polygenes had additive effect, dominant effect and sometimes had epistatic effect. The major gene was partial dominant or complete dominant and the two pairs of major genes had the same effects in one hybrid combination and had the different effects in other combinations. The polygenes were partial dominant, complete dominant or superdominance in all combinations. Heritabilities of major gene were greater than those of polygenes in the three segregating generations. The estimate values of heritability of major gene and polygenes were relatively high. These results suggested that the inheritance of B. oleracea seed colour was a considerably complex system, and the effects of seed colour genes were quite different in different hybrid combinations and belonged to different genetic models.
(3) The performance of B. oleracea seed colour could be affected by enviromental factors in a certain degree.
2. The effects of agronomic measures and genotype×environment interaction on B. oleracea seed colour
B. oleracea var. acephala, B. oleracea var. capitata and B. oleracea var. alboglabra with different seed colours were used as materials and split-split plot design and randomized block design were adopted to investigate the effects of different agronomic measures, including the amount of nitrogen fertilizer, planting density, sowing and harvest time, and genotype×environment interaction on seed colour, as well as the performance stability of seed colours to the environmental changes in B. oleracea. The main conclusions are as followings:
(1) The effects of the amount of nitrogen fertilizer, sowing and harvest time on seed colour varied with genotypes in B. oleracea. No obvious effects on the seed colour of black-seeded genotypes and very significant influences on the yellow-seeded genotypes were found. The seed colour of yellow-seeded genotypes would gradually become lighter with the amount increase of nitrogen fertilizer. The early or late sowing time would make the seed colour of yellow-seeded genotypes dark. The too late harvest time could extremely significantly reduce the seed yellow degree of yellow-seeded genotypes. Planting density had no apparent effects on B. oleracea seed colour.
(2) The interaction of genotype×environment significantly affected B. oleracea seed colour and different genotypes had different reactions to the environment changes, especially in the yellow-seeded genotypes.
3. The dynamic changes of seed coat pigment content during seed development process in B. oleracea
Three pairs of B. oleracea var. acephala yellow-and black-seeded lines with the same genetic background from different sources were used to analyze the dynamic changes of main seed coat pigment content (chlorophyll, carotenoid, anthocyanidin, melanin and flavonoid) during the seed development process and the differences of seed coat pigment content in mature seed. The main conclusions are as followings:
(1) In the seed development process of B. oleracea, the contents of chlorophyll, anthocyanidin, melanin and flavonoid except carotenoid showed significant differences in seed coat with different colour and all the five pigment contents varied significantly at different development stages. The chlorophyll, anthocyanidin and flavonoid presented a change trend from increasing to decreasing. The carotenoid contents of black seed coat monotonicly increased with the seed development, whereas those of yellow seed coat decreased at the early stage and then increased. Melanin contents increased sharply from40days after flowering and the differences of melanin content enlarged rapidly between black and yellow seed coat. The carotenoid contents of yellow and black seed coat were almost at the same level in mature seed, whereas the contents of chlorophyll, anthocyanidin, melanin and flavonoid of black seed coat were extremely significantly or significantly higher than those of yellow seed coat which had the same genetic background. The different change tendencies of the five pigment contents existed between yellow and black seed coat in the process of seed development.
(2) The contents of chlorophyll, anthocyanidin and flavonoid of seed coat would continue to decline in B. oleracea, whereas the contents of carotenoid and melanin continue to rise after harvest and threshing. The five pigment contents of black seed coat were significantly higher than those of yellow seed coat with the same genetic background in the air-drying mature seed. Comparing with yellow seed coat, the anthocyanidin content was the highest in black seed coat, the contents of melanin and chlorophyll were next, and the contents of carotenoid and flavonoid were the last. Therefore, in air-drying mature seed coat of B. oleracea, the main pigment causing the differences of seed colour was anthocyanidin, next melanin and chlorophyll, and then carotenoid and flavonoid.
4. Comparative analysis of seed colour inheritance in B. oleracea and B. napus
The means of literature research were used to compare and analyze the genetic characteristics of seed colour of B. oleracea and B. napus, the effects of major agronomic measures and genotype×environment interaction on seed colour, the stability of seed colour to the environmental changes, the dynamic changes of main pigment content of seed coat during seed development process and the differences of pigment content in mature seed coat, etc. The main conclusions are as followings:
(1) The hereditary characteristics of B. oleracea seed colour were similar to B. napus and there were no obvious differences in the dominant or recessive gene of yellow seed trait, the number and effect of seed colour gene, as well as maternal effect and heritability, etc.
(2) Except planting density, the effects of major agronomic measures, such as sowing and harvest time and the amount of nitrogen fertilizer, etc, on seed colour were relatively similar between B. oleracea and B. napus. The genotype×environment interaction effect on B. oleracea seed colour and the environmental stability of B. oleracea seed colour were similar to B. napus.
(3) During seed development process, in addition to a slight difference of peak date of anthocyanidin content, the dynamic changes of chlorophyll, carotenoid, melanin and flavonoids were no significant differences between B. oleracea and B. napus seed coat. Between yellow and black mature seed coat, the difference of chlorophyll content in B. oleracea was different with B. napus, the rest four pigments were the same. The largest difference between yellow and black mature seed coat was anthocyanidin content in B. oleracea, followed by melanin and chlorophyll, however they were melanin and anthocyanidin in B. napus.
(4) It suggested that quantitative genetic characteristics of yellow seed trait of B. napus was likely to be resulted from the genes on the C genome come from B. oleracea.
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