若干十字花科植物小孢子培养和植株再生及四倍体新种质创制
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摘要
倍性育种在十字花科植物育种中具有很重要的作用。小孢子培养可短时间内获得纯合、稳定的育种材料,从而加快了选择进程,缩短了育种年限,并且还能分离得到隐形性状,在创制优良自交系和提高育种效率方面具有重要作用。多倍体植株的“巨大性”有利于提高植物中有机成分含量和植株生物产量,在蔬菜及具有特殊用途的作物育种中具有重要价值。
本研究以若干十字花科植物(油菜、甘蓝、青花菜、羽衣甘蓝、萝卜)为试材,对影响小孢子培养和植株再生的因素进行研究；人工诱导创制富含花青素四倍体红皮红心萝卜新种质,并对其中花青素分布、不同部位含量等方面进行了研究,获得主要研究成果如下：
1、对上述十字花科植物小孢子出胚率和胚状体再生的影响因素进行了研究,结果表明,基因型是影响小孢子胚胎发生的关键因素；4℃低温预处理1d并结合32.5℃热激1d,能显著提高供试基因型小孢子出胚率；在固体分化培养基中添加1%-1.2%琼脂有利于小孢子胚萌发和植株再生；胚在液体培养基中滞留时间以25d左右为最好。
2、采用DAPI染色法对几个甘蓝类植物(甘蓝、青花菜、羽衣甘蓝)的小孢子发育时期与花蕾大小、形态的关系进行观察,并与易出胚油菜品种‘浙双758’一起进行游离小孢子培养。结果表明,不同供试基因型处于单核晚期的花蕾大小之间存在差异；P/A(花瓣/花药)≈1可以作为几个甘蓝类植物选择合适花蕾的标准。不同供试基因型与油菜小孢子共培养的胚胎发生频率和植株再生率均高于对照,然而不同基因型间胚胎发生率存在差异；小孢子再生植株群体中,甘蓝类植物小孢子再生植株形态特征与油菜相比差异显著,可以直接进行区别。结果发现,共培养无论是在小孢子胚胎发生率和胚胎质量,还是再生植株率,均显著优于未共培养对照。
3、采用DAPI荧光染色法可清楚地观察到小孢子发育时期。甘蓝花蕾平均长为3.0-4.5mm、花瓣/药为0.85-1.20时,处于单核晚期至双核早期的小孢子所占比例最多；青花菜小孢子处于单核晚期至双核早期时,其花蕾平均长为2.0-3.5mm、花瓣/药为0.85-1.20；羽衣甘蓝和萝卜花蕾平均长分别为为3.5～4.5mm、3.0-4.0mm、花瓣/药分别为0.9-1.2、0.9-1.15时最适于进行小孢子培养实验。显微观察发现,小孢子受到高温刺激后,体积开始膨大,孢子壁逐渐分解,大部分小孢子发生对称分裂。培养15d左右时,发育成肉眼可见的胚状体,25d左右时,小孢子发育成胚状体且适宜于再生成植株。小孢子感受到热激后,电镜下可观察到小孢子细胞中液泡部分或完全消失,细胞核移到细胞中央,细胞进入到第一次分裂前的准备阶段；细胞器大量聚集在细胞质中,细胞核周围存在着丰富的淀粉粒；小孢子发生第一次分裂后形成两个大小、形态相似的细胞,两个细胞核着色较浅,细胞器均匀分布在细胞质中,两个细胞相邻但没有细胞壁包被。
4、小孢子再生植株群体中,除单倍体外,还存在二倍体和多倍体植株,采用流式细胞仪可快速鉴定出群体中倍性水平。采用200mg·L-1水仙碱处理单倍体植株20h的诱导效果最好,3个供试基因型植株的存活率和二倍体诱导率均优于其它处理。不同倍性间植株气孔大小和形态以及保卫细胞叶绿体数存在差异,采用FDA荧光染色和碘-碘化钾染色法可清楚地观察到不同倍性间的差别。发现单倍体植株气孔长度约为20μm,二倍体约为25μm,而四倍体大于33μm；单倍体保卫细胞叶绿体数为7个左右,二倍体约13个,而四倍体最多超过23个,差异显著。
5、采用不同浓度秋水仙碱(0.1,0.15,0.2g·L-1不同处理时间(0,24,48,72h)组合对‘胭脂3号’萝卜子叶期生长点进行处理；先根据形态学特征和气孔大小形态对处理群体进行初步鉴定,得到的疑似变异植株再采用流式细胞仪和染色体计数法进行最终倍性确定；并对得到的四倍体和二倍体红皮红心萝卜的形态学特征和总花青素含量进行比较。结果表明,秋水仙碱浓度越高、处理时间越长,植株死亡率越高；0.2g·L-1仙碱处理72h的植株变异率最高,为62.0%；0.2g·L-1水仙碱处理48h得到的四倍体诱导率最高,为11.3%,共有37株加倍为四倍体。四倍体和二倍体植株在叶片和花形态、植株长势、花粉粒和气孔形态以及保卫细胞叶绿体数目等表现差异；四倍体红皮红心萝卜花青素含量比二倍体增加11.8%,差异显著。
6、对红心萝卜生长过程中花青素合成规律及与PAL酶活性、可溶性糖含量相关性进行分析,结果表明,花青素在种子萌发时即开始合成,合成顺序是从上而下的,且在下胚轴中的分布是不均匀的；在肉质根中,花青素在根皮中的含量高于根肉,根上部含量高于下部,与在下胚轴中分布方式一致。光照条件下子叶和下胚轴中的花青素、PAL酶活性、可溶性糖含量均比黑暗条件下的高。光照条件下下胚轴和田间生长过程中肉质根中花青素含量呈上升趋势；而光照条件下子叶和黑暗条件下的子叶和下胚轴中的花青素含量均呈先升高后降低的变化趋势。PAL酶活性、可溶性糖含量在发芽和田间生长过程中的变化趋势不同。叶片中叶绿素含量在萝卜生长过程中呈现波动增加的变化趋势。肉质根皮中花青素含量最高,为25.03mg/1OOg,其次为肉质部,根头和叶柄中含量较少。PAL酶活性、可溶性糖含量均与花青素含量呈正相关,相关系数分别为0.607和0.678；叶绿素与花青素也呈正相关,相关系数为0.496。
Ploidy breeding is very important in Cruciferous species breeding. The isolated microspore culture can obtain pure and stable materials in short time, accelerate breeding process, shorten breeding times and get recessive characters. It is useful in creating excellent inbred lines and promoting breeding efficient. The content of biological component and biomass yield in polyploidy plants is promoted than in diploid. It is very important in vegetables and other special useful crops.
In the present paper, several Cruciferous species, such as cabbage, broccoli, kale and radish, were used to study the factors on microspore embryogenesis and plantlet regeneration. The tetraploidy radish, rich in red-pigment, was induced by colchicine treatment, and anthocyanin distribution and content in different parts were studied. Main results are listed as follows:
1. The factors that affect embryogenesis and plant regeneration efficiency of several cultivars of Cruciferous species (cabbage, broccoli, kale and radish) were studied. The results showed that genotype was key factor that influenced microspore embryogenesis;4℃cold temperature pre-treated for1d combining32.5℃treated for1d could improve embryogenesis efficiency significantly. In solid differentiation medium,1-2%concentration of agar was suitable to microspore embryo germination and plantlet regeneration. The embryos cultured in NLN-13liquid medium about25d were best for germination.
2. The floewer bud with microspore at late uninucleate stage was investigated by DAPI staining, and microspores of different B. oleracea species were co-cultured with that of B. napus (good responsive to microspore embryogenesis). The results showed that the size of buds containing late uninucleate microspores was different in various genotypes. The ratio of P/A (petal/anther) around1could be used as an index to select buds of B. oleracea species. Co-cultured with microspores of B. napus, cv. Zheshuang758, different genotypes of B. oleracea species exhibited improved frequency of embryo production and plant regeneration than those of non-co-culture controls. The efficiency of embryogenesis was different among different B. oleracea species. Microspore-derived plants of different cultivars of B. oleracea were discriminated from B. napus according to the morphological characters of plant. The results indicated that B. oleracea produced higher number of microspore-derived plants in co-culture treatments than controls.
3. DAPI staining was used to observe microspore developmental stage and found that good results were observed from microspore at late uninucleate to early binucleate stage, with the flower buds average length of3.0-5.0mm and P/A of0.8-1.2for cabbage;2.5-4.0mm and P/A of0.8-1.25for broccoli;3.0-5.0mm and P/A of0.8-1.2for kale, and3.0-4.0mm and P/A of0.9-1.2for radish. When microspores experienced heat shock, the volume began to expand, exine broke down gradually and many microspores entered into symmetric division. About15d after culture, macroscopic embryos showed up and developed into embryoids25d later which were suitable to transfer into solid differentiation medium. After heat shock, the vacuoles in microspore vanish, nucleus moved to center and microspores prepared for the first division. Simultaneously, lots of organelles gathered in cytoplasm and nucleus around abundant of starch grains. After first division, two similar cells were developed and the two nucleuses were light in color, and organelles spread in cytoplasm uniformly. The two cells closed each other but no cell wall envelope.
4. In microspore-derived plant population, the ploidy level was mixed. Besides haploids, there were also diploids and other polyploidy plants. Flow cytometry analysis could be used to identify the ploidy level rapidly. Various concentrations of colchicine for different durations were used to treat haploid plants. The results indicated that the combination of200mg-L"1colchicine for20h was better than other treatments in term of plant survival and diploid inducement efficiency in three rested genotypes. The differences existed in stomatal size and morphologies, and guard cell chloroplasts in different ploidy level. The methods of FDA and I-KI staining could be used to observe the differences clearly. We found that stomatal length of haploid was approximately20μm, and25μm for diploid, while more than33μm for tetraploid. The number of guard cell chloroplasts in haploid was about7, and13for diploid and more than23for tetraploid. The difference of number of guard cell chloroplasts was significant in different ploidy levels.
5. The apical tips of cotyledon-stage seedlings of a unique red-core radish cultivar "Yanzhi" were treated with various concentrations of colchicines (0.1,0.15,0.2g-L-1) for different durations (0,24,48,72h). The morphological characteristics and stomatal size were used to pre-screen putative polyploids one month later after treatment, and then ploidy level was confirmed by using the flow cytometry and chromosome counting. Morphologies and total anthocyanin content were compared between tetraploid and diploid plants of red-core radish. The results indicated that high concentration of colchicine treatments resulted in high mortality rate. The highest percent of variation plants was62.0%when treated with0.2g·L-1colchicine for72h, and the highest tetraploid induction efficiency was11.3%when seedlings treated with0.2g·L-1colchicine for48h where37plants of these putative tetraploids were indeed doubled. The tetraploid and diploid plantlets differed significantly in leaves and flower morphologies, plant vitality and also the stomatal and pollen grain sizes and guard cell chloroplasts. The anthocyanin content in tetraploid plants was173.6mg/100g FW,11.8%higher than that of diploid plants significantly.
6、The synthesis of anthocyanin and their correlation to phenylalanine ammonia-lyase (PAL) activity, soluble sugar and chlorophyll contents were investigated in the present experiment. The results showed that the anthocyanin was synthesized during germination. The configuration order of anthocyanin was from top to bottom in hypocotyls and distributed in a non-uniform manner. The anthocyanin content in root peels was higher than that in flesh and was higher in aboveground parts as compared to underground parts. The contents of pigment, soluble sugar, and PAL activity were higher in light than in the dark. The changes of anthocyanin content of hypocotyls in roots under the light were raised. The trend of cotyledons in the light and cotyledons and hypocotyls in the dark increased initially and then reduced. Light influenced the trends of PAL activity and soluble sugar content during germination and growth in the field in a different mode. The chlorophyll content in leaves was fluctuated during growth period and the trend was on the rise, but almost unchanged in the petioles. In roots, the content of pigment in the peels was the highest (25.03mg/100g), followed by flesh and root apex, and least in petioles. The correlation of pigment to PAL activity and soluble sugar content was positive, but the correlation coefficient was different (at0.607and0.678, respectively). The relevance of chlorophyll to pigment was also positive with the correlation coefficient of0.496.
本研究以若干十字花科植物(油菜、甘蓝、青花菜、羽衣甘蓝、萝卜)为试材,对影响小孢子培养和植株再生的因素进行研究；人工诱导创制富含花青素四倍体红皮红心萝卜新种质,并对其中花青素分布、不同部位含量等方面进行了研究,获得主要研究成果如下：
1、对上述十字花科植物小孢子出胚率和胚状体再生的影响因素进行了研究,结果表明,基因型是影响小孢子胚胎发生的关键因素；4℃低温预处理1d并结合32.5℃热激1d,能显著提高供试基因型小孢子出胚率；在固体分化培养基中添加1%-1.2%琼脂有利于小孢子胚萌发和植株再生；胚在液体培养基中滞留时间以25d左右为最好。
2、采用DAPI染色法对几个甘蓝类植物(甘蓝、青花菜、羽衣甘蓝)的小孢子发育时期与花蕾大小、形态的关系进行观察,并与易出胚油菜品种‘浙双758’一起进行游离小孢子培养。结果表明,不同供试基因型处于单核晚期的花蕾大小之间存在差异；P/A(花瓣/花药)≈1可以作为几个甘蓝类植物选择合适花蕾的标准。不同供试基因型与油菜小孢子共培养的胚胎发生频率和植株再生率均高于对照,然而不同基因型间胚胎发生率存在差异；小孢子再生植株群体中,甘蓝类植物小孢子再生植株形态特征与油菜相比差异显著,可以直接进行区别。结果发现,共培养无论是在小孢子胚胎发生率和胚胎质量,还是再生植株率,均显著优于未共培养对照。
3、采用DAPI荧光染色法可清楚地观察到小孢子发育时期。甘蓝花蕾平均长为3.0-4.5mm、花瓣/药为0.85-1.20时,处于单核晚期至双核早期的小孢子所占比例最多；青花菜小孢子处于单核晚期至双核早期时,其花蕾平均长为2.0-3.5mm、花瓣/药为0.85-1.20；羽衣甘蓝和萝卜花蕾平均长分别为为3.5～4.5mm、3.0-4.0mm、花瓣/药分别为0.9-1.2、0.9-1.15时最适于进行小孢子培养实验。显微观察发现,小孢子受到高温刺激后,体积开始膨大,孢子壁逐渐分解,大部分小孢子发生对称分裂。培养15d左右时,发育成肉眼可见的胚状体,25d左右时,小孢子发育成胚状体且适宜于再生成植株。小孢子感受到热激后,电镜下可观察到小孢子细胞中液泡部分或完全消失,细胞核移到细胞中央,细胞进入到第一次分裂前的准备阶段；细胞器大量聚集在细胞质中,细胞核周围存在着丰富的淀粉粒；小孢子发生第一次分裂后形成两个大小、形态相似的细胞,两个细胞核着色较浅,细胞器均匀分布在细胞质中,两个细胞相邻但没有细胞壁包被。
4、小孢子再生植株群体中,除单倍体外,还存在二倍体和多倍体植株,采用流式细胞仪可快速鉴定出群体中倍性水平。采用200mg·L-1水仙碱处理单倍体植株20h的诱导效果最好,3个供试基因型植株的存活率和二倍体诱导率均优于其它处理。不同倍性间植株气孔大小和形态以及保卫细胞叶绿体数存在差异,采用FDA荧光染色和碘-碘化钾染色法可清楚地观察到不同倍性间的差别。发现单倍体植株气孔长度约为20μm,二倍体约为25μm,而四倍体大于33μm；单倍体保卫细胞叶绿体数为7个左右,二倍体约13个,而四倍体最多超过23个,差异显著。
5、采用不同浓度秋水仙碱(0.1,0.15,0.2g·L-1不同处理时间(0,24,48,72h)组合对‘胭脂3号’萝卜子叶期生长点进行处理；先根据形态学特征和气孔大小形态对处理群体进行初步鉴定,得到的疑似变异植株再采用流式细胞仪和染色体计数法进行最终倍性确定；并对得到的四倍体和二倍体红皮红心萝卜的形态学特征和总花青素含量进行比较。结果表明,秋水仙碱浓度越高、处理时间越长,植株死亡率越高；0.2g·L-1仙碱处理72h的植株变异率最高,为62.0%；0.2g·L-1水仙碱处理48h得到的四倍体诱导率最高,为11.3%,共有37株加倍为四倍体。四倍体和二倍体植株在叶片和花形态、植株长势、花粉粒和气孔形态以及保卫细胞叶绿体数目等表现差异；四倍体红皮红心萝卜花青素含量比二倍体增加11.8%,差异显著。
6、对红心萝卜生长过程中花青素合成规律及与PAL酶活性、可溶性糖含量相关性进行分析,结果表明,花青素在种子萌发时即开始合成,合成顺序是从上而下的,且在下胚轴中的分布是不均匀的；在肉质根中,花青素在根皮中的含量高于根肉,根上部含量高于下部,与在下胚轴中分布方式一致。光照条件下子叶和下胚轴中的花青素、PAL酶活性、可溶性糖含量均比黑暗条件下的高。光照条件下下胚轴和田间生长过程中肉质根中花青素含量呈上升趋势；而光照条件下子叶和黑暗条件下的子叶和下胚轴中的花青素含量均呈先升高后降低的变化趋势。PAL酶活性、可溶性糖含量在发芽和田间生长过程中的变化趋势不同。叶片中叶绿素含量在萝卜生长过程中呈现波动增加的变化趋势。肉质根皮中花青素含量最高,为25.03mg/1OOg,其次为肉质部,根头和叶柄中含量较少。PAL酶活性、可溶性糖含量均与花青素含量呈正相关,相关系数分别为0.607和0.678；叶绿素与花青素也呈正相关,相关系数为0.496。
Ploidy breeding is very important in Cruciferous species breeding. The isolated microspore culture can obtain pure and stable materials in short time, accelerate breeding process, shorten breeding times and get recessive characters. It is useful in creating excellent inbred lines and promoting breeding efficient. The content of biological component and biomass yield in polyploidy plants is promoted than in diploid. It is very important in vegetables and other special useful crops.
In the present paper, several Cruciferous species, such as cabbage, broccoli, kale and radish, were used to study the factors on microspore embryogenesis and plantlet regeneration. The tetraploidy radish, rich in red-pigment, was induced by colchicine treatment, and anthocyanin distribution and content in different parts were studied. Main results are listed as follows:
1. The factors that affect embryogenesis and plant regeneration efficiency of several cultivars of Cruciferous species (cabbage, broccoli, kale and radish) were studied. The results showed that genotype was key factor that influenced microspore embryogenesis;4℃cold temperature pre-treated for1d combining32.5℃treated for1d could improve embryogenesis efficiency significantly. In solid differentiation medium,1-2%concentration of agar was suitable to microspore embryo germination and plantlet regeneration. The embryos cultured in NLN-13liquid medium about25d were best for germination.
2. The floewer bud with microspore at late uninucleate stage was investigated by DAPI staining, and microspores of different B. oleracea species were co-cultured with that of B. napus (good responsive to microspore embryogenesis). The results showed that the size of buds containing late uninucleate microspores was different in various genotypes. The ratio of P/A (petal/anther) around1could be used as an index to select buds of B. oleracea species. Co-cultured with microspores of B. napus, cv. Zheshuang758, different genotypes of B. oleracea species exhibited improved frequency of embryo production and plant regeneration than those of non-co-culture controls. The efficiency of embryogenesis was different among different B. oleracea species. Microspore-derived plants of different cultivars of B. oleracea were discriminated from B. napus according to the morphological characters of plant. The results indicated that B. oleracea produced higher number of microspore-derived plants in co-culture treatments than controls.
3. DAPI staining was used to observe microspore developmental stage and found that good results were observed from microspore at late uninucleate to early binucleate stage, with the flower buds average length of3.0-5.0mm and P/A of0.8-1.2for cabbage;2.5-4.0mm and P/A of0.8-1.25for broccoli;3.0-5.0mm and P/A of0.8-1.2for kale, and3.0-4.0mm and P/A of0.9-1.2for radish. When microspores experienced heat shock, the volume began to expand, exine broke down gradually and many microspores entered into symmetric division. About15d after culture, macroscopic embryos showed up and developed into embryoids25d later which were suitable to transfer into solid differentiation medium. After heat shock, the vacuoles in microspore vanish, nucleus moved to center and microspores prepared for the first division. Simultaneously, lots of organelles gathered in cytoplasm and nucleus around abundant of starch grains. After first division, two similar cells were developed and the two nucleuses were light in color, and organelles spread in cytoplasm uniformly. The two cells closed each other but no cell wall envelope.
4. In microspore-derived plant population, the ploidy level was mixed. Besides haploids, there were also diploids and other polyploidy plants. Flow cytometry analysis could be used to identify the ploidy level rapidly. Various concentrations of colchicine for different durations were used to treat haploid plants. The results indicated that the combination of200mg-L"1colchicine for20h was better than other treatments in term of plant survival and diploid inducement efficiency in three rested genotypes. The differences existed in stomatal size and morphologies, and guard cell chloroplasts in different ploidy level. The methods of FDA and I-KI staining could be used to observe the differences clearly. We found that stomatal length of haploid was approximately20μm, and25μm for diploid, while more than33μm for tetraploid. The number of guard cell chloroplasts in haploid was about7, and13for diploid and more than23for tetraploid. The difference of number of guard cell chloroplasts was significant in different ploidy levels.
5. The apical tips of cotyledon-stage seedlings of a unique red-core radish cultivar "Yanzhi" were treated with various concentrations of colchicines (0.1,0.15,0.2g-L-1) for different durations (0,24,48,72h). The morphological characteristics and stomatal size were used to pre-screen putative polyploids one month later after treatment, and then ploidy level was confirmed by using the flow cytometry and chromosome counting. Morphologies and total anthocyanin content were compared between tetraploid and diploid plants of red-core radish. The results indicated that high concentration of colchicine treatments resulted in high mortality rate. The highest percent of variation plants was62.0%when treated with0.2g·L-1colchicine for72h, and the highest tetraploid induction efficiency was11.3%when seedlings treated with0.2g·L-1colchicine for48h where37plants of these putative tetraploids were indeed doubled. The tetraploid and diploid plantlets differed significantly in leaves and flower morphologies, plant vitality and also the stomatal and pollen grain sizes and guard cell chloroplasts. The anthocyanin content in tetraploid plants was173.6mg/100g FW,11.8%higher than that of diploid plants significantly.
6、The synthesis of anthocyanin and their correlation to phenylalanine ammonia-lyase (PAL) activity, soluble sugar and chlorophyll contents were investigated in the present experiment. The results showed that the anthocyanin was synthesized during germination. The configuration order of anthocyanin was from top to bottom in hypocotyls and distributed in a non-uniform manner. The anthocyanin content in root peels was higher than that in flesh and was higher in aboveground parts as compared to underground parts. The contents of pigment, soluble sugar, and PAL activity were higher in light than in the dark. The changes of anthocyanin content of hypocotyls in roots under the light were raised. The trend of cotyledons in the light and cotyledons and hypocotyls in the dark increased initially and then reduced. Light influenced the trends of PAL activity and soluble sugar content during germination and growth in the field in a different mode. The chlorophyll content in leaves was fluctuated during growth period and the trend was on the rise, but almost unchanged in the petioles. In roots, the content of pigment in the peels was the highest (25.03mg/100g), followed by flesh and root apex, and least in petioles. The correlation of pigment to PAL activity and soluble sugar content was positive, but the correlation coefficient was different (at0.607and0.678, respectively). The relevance of chlorophyll to pigment was also positive with the correlation coefficient of0.496.
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