红三叶新品系种质鉴定及无性繁殖特性的研究
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摘要
红三叶新品系(Trifolium pratense L.)是内蒙古农业大学以内蒙古牙克石地区自然分布的红三叶种质为材料,经过多年的栽培驯化和系统选育,而培育出的抗寒新种质。本文对红三叶新品系以及部分国内外种质的形态学特征、染色体、SSR标记以及无性繁殖做了较为系统的鉴定评价和研究,并首次对内蒙古牙克石自然分布的红三叶遗传起源进行了探讨,为我国红三叶优异种质的利用和创新奠定了基础。研究获得以下主要结果：
1、70份红三叶材料中早花型、中间型和晚花型,分别占22.86%、68.57%和8.57%。根据第二年生育期长短可分为早熟型、中熟型和晚熟型,生育期分别64d,71.96d和84.17d。红三叶新品系属晚花型,生育期74d,中熟型,平均日增长速率为0.44cm/d,接近红三叶群体平均水平。试验材料在呼和浩特越冬率变化范围为0%～88.89%,红三叶新品系越冬率88.89%,抗寒性突出。
2、形态特征研究表明,红三叶叶斑强度、分枝数、花序数/茎、种子颜色、第三茎节长、二级侧枝数、叶面积、自然高度等形态特征变异较大。遗传多样性由大到小依次为枝形>叶形>种子>花部>根部。冠幅、自然高度和分枝数是划分红三叶形态类型的主要指标。红三叶新品系根系以黄褐色为主,多为绿色茎,半直立,株丛大小为红三叶群体中等水平,叶斑以中斑和无斑为主,叶面积较大,叶片形状较为宽短。花色主要为紫红色,浅紫红色次之,浅粉色出现比例约为10%,偶见白花的变异。种子颜色相对较浅,黄色种子数量较多,种子大小居中,千粒重偏小,红三叶新品系与29,35,3,41,5,19,2,67号等材料形态特征比较相似。
3、43份红三叶均为二倍体,染色体数目为2n=14,基数为7,共检测出m、sm、 st和t4种染色体类型,其中6份材料出现随体,有1A、2A、1B、2B和3B5种核型,8种核型公式。红三叶新品系核型公式为2n=2x=14=8m+6sm(2SAT),染色体内部不对称性大于染色体之间的不对称性,不对称性在红三叶群体中相对较高。根据Langlet染色体形态划分标准,红三叶新品系染色体类型属于相对原始的T型。
4、32份红三叶材料,8对引物扩增共获得多态性条带219条,多态性百分率为100%,有效等位基因数Ne为1.1636。Nei多样性指数H为0.1229,Shannon信息指数Ⅰ为0.2241,材料间的总变异Ht为0.1218,种群间遗传变异Hs0.0387,遗传分化系数Gst为0.3179,基因流Nm1.0727,31.79%的遗传变异存在于种群之间,68.21%的存在于种群内。本试验获得红三叶新品系的指纹图谱,可有效区分2份国内已登记红三叶品种。红三叶新品系Nei多样性指数0.0665,Shannon信息指数为0.1065,均低于平均水平,种群多样性相对较低。遗传距离表明红三叶新品系亲缘最近的3份材料依次为来自俄罗斯的60号>西班牙的22号>俄罗斯的53号。
5、红三叶新品系分株繁殖适宜切块6-7个,最高增殖系数为6.38,分枝期整枝扦插效率较高,最高增殖系数为6.40。组织培养研究表明：红三叶新品系种子消毒的最佳方法为75%酒精预处理,0.1%HgCl2消毒10min；最适无菌苗培养基为不添加激素的1/4MS+1.5%蔗糖；子叶、下胚轴、叶柄、叶片及根段在MS+0.5mg·L-16-BA+1～2mg·L-12,4-D培养基上都极易诱导出愈伤组织；最佳继代培养基为MS+0.5mg.L-16-BA+1.0mg·L-12,4-D;最佳分化培养基为MS+0.75mg·L-16-BA+0.5mg·L-1NAA,最高分化率为16.67%；茎芽增殖最佳培养基为MS+0.5mg·L-16-BA+1.0mg·L-1IBA;最佳生根条件为50mg·L-1浓度的IBA中浸泡1h后植入含0.5mg·L-1IBA的MS中,生根率达80%。1mg·L-1BA+0.5mg·L-1NAA激素配比适合红三叶茎尖和腋芽的萌发。
6、综合分析,初步推断内蒙古牙克石自然分布的红三叶可能起源于俄罗斯西部,属于欧洲血统。
The new line of red clover (Trifolium pratense L.) is an excellent cold resistant material suitable for alpine region in China, which filled the blank of red clover breeding in Inner Mongolia. Red clover new line and some red clover materials were studied by morphological features, chromosome, SSR markers, and asexual reproduction. The genetic origin of red clover which is natural distributed in Yakeshi in Inner Mongolia was discussed at first time in this paper. There is a great significance of the use and innovation of elite germplasm in red clover. Obtained the following results:
1、70materials can be divided into the early flowering, the middle flowering and late flowering, respectively, accounted for22.86%,68.57%and8.57%. According to growth period in second year, red clover can be divided into the early, medium and late maturity groups, and growth period were64d,71.96d and84.17d, respectively. New lines of red clover is a late flowering type. Growth period was74d belongs to the medium. Average growth rates in daily was0.44cm/d which close to the average level of red clover groups. Winter survival rates were from0%to88.89%in70materials at Hohhot. The new line of red clover winter rate was88.89%, and cold resistance was outstanding.
2、The studies of morphological characteristics have shown that red clover organs have great genetic variation. The coefficient of variation on leaf mark, branches, inflorescences per stem, seed color, internode length, lateral branch number, leaf area, natural height were relatively larger. The order of genetic diversity from the largest to the smallest was branches> leaves> seeds> flowers> roots. The crown width, height and branches were main indicators to divide morphological types of red clover. New lines of red clover had tan roots, and usually green stem which was semi-erect. Strains clump size was the middle in the red clover groups. Leaf mark were middle or no mark mainly. Leaf area was bigger relatively, and leaf shape was wider and shorter. The main flower color was purple,then light purple and pink, white occasionally. Seed color was relatively shallow, ana a large number of seeds was yellow. Seed size was middle, and1000-grain weight was smaller relative to the red clover groups. The new line of red clover was similar to the number of29,35,3,41,5,19,2,67in morphological characteristics.
3、Chromosome number of43red clover was2n-2x=14, x=7. Chromosome type included m, sm, st, and t, six of which had satellite. There were5kinds karyotype (1A,2A,1B,2B and3B)and8kinds of karyotype formula. The karyotype formula of new line of red clover was2n=2x=14=8m+6sm(2SAT). The asymmetry of interchromosomal is greater than intrachromosomal, and was relatively high in the red clover populations. According to the the Langlet's chromosome morphology criteria for the classification, the new line of red clover chromosome type is the T-type, primitive relatively.
4、The analysis of SSR marker showed that using8pairs of primers in32materials obtain219polymorphic bands totally. The polymorphism percentage was100%, and the number of effective alleles was1.1636. The Nei'diversity index, shannon information index, total variance, variance between the materials, coefficient of genetic differentiation and gene flow were0.1229,0.2241,0.1218,0.0387,0.3179respectively.31.79%genetic variation of the red clover exists in external populations,68.21%was in internal populations. The fingerprints of the red clover new line was obtained, which can effectively distinguish from two registered varieties of red clover in China. The Nei diversity index of red clover new line was0.0665, and Shannon's information index was0.1065, lower than the average population diversity level. SSR genetic distance showed that the new line of red clover was similar to the number of60,22,53, which were from Russia, Spain and Russia respectively.
5、The best ramets per plant was6-7in division propagation, and the highest proliferation coefficient was6.38; The highest proliferation coefficient of cutting can up to6.40using entire branches in branching period.The study of tissue culture showed:the best method of seeds sterilization for red clover is the75% ethyl alcohol treatment followed by disinfection with0.1%HgCl2for10min.1/4MS medium with1.5%sucrose without hormones was most effective for seed culture. Cotyledons, hypocotyl, petiole, leaf and root callus were easy to be induced in medium of MS containing0.5mg·L'16-BA and1-2mg·L-12,4-D. The best callus subculture medium was MS containing0.5mg·L-116-BA and1.0mg·L-12,4-D. The best medium for differentiation was MS plus0.75mg·L-16-BA and0.5mg·L-1NAA, the differentiation rate can reach up to16.67%. Corm-bud could be multiplicated greatly in MS medium plus0.5mg·L-16-BA and1.0mg·L-1IBA. Regenerated plants were immersed in50mg·L-1IBA for1h, then vaccinated in MS medium plus IBA0.5mg·L-1,the rooting rate was80%.1mg·L-16-BA+0.5mg·L-1NAA hormone combination was suitable for germination of shoot tips and axillary buds in red clover.
6、We concluded that red clover that distributed at Yakeshi in Inner Mongolia is likely to come from western Russia and was a European descent.
1、70份红三叶材料中早花型、中间型和晚花型,分别占22.86%、68.57%和8.57%。根据第二年生育期长短可分为早熟型、中熟型和晚熟型,生育期分别64d,71.96d和84.17d。红三叶新品系属晚花型,生育期74d,中熟型,平均日增长速率为0.44cm/d,接近红三叶群体平均水平。试验材料在呼和浩特越冬率变化范围为0%～88.89%,红三叶新品系越冬率88.89%,抗寒性突出。
2、形态特征研究表明,红三叶叶斑强度、分枝数、花序数/茎、种子颜色、第三茎节长、二级侧枝数、叶面积、自然高度等形态特征变异较大。遗传多样性由大到小依次为枝形>叶形>种子>花部>根部。冠幅、自然高度和分枝数是划分红三叶形态类型的主要指标。红三叶新品系根系以黄褐色为主,多为绿色茎,半直立,株丛大小为红三叶群体中等水平,叶斑以中斑和无斑为主,叶面积较大,叶片形状较为宽短。花色主要为紫红色,浅紫红色次之,浅粉色出现比例约为10%,偶见白花的变异。种子颜色相对较浅,黄色种子数量较多,种子大小居中,千粒重偏小,红三叶新品系与29,35,3,41,5,19,2,67号等材料形态特征比较相似。
3、43份红三叶均为二倍体,染色体数目为2n=14,基数为7,共检测出m、sm、 st和t4种染色体类型,其中6份材料出现随体,有1A、2A、1B、2B和3B5种核型,8种核型公式。红三叶新品系核型公式为2n=2x=14=8m+6sm(2SAT),染色体内部不对称性大于染色体之间的不对称性,不对称性在红三叶群体中相对较高。根据Langlet染色体形态划分标准,红三叶新品系染色体类型属于相对原始的T型。
4、32份红三叶材料,8对引物扩增共获得多态性条带219条,多态性百分率为100%,有效等位基因数Ne为1.1636。Nei多样性指数H为0.1229,Shannon信息指数Ⅰ为0.2241,材料间的总变异Ht为0.1218,种群间遗传变异Hs0.0387,遗传分化系数Gst为0.3179,基因流Nm1.0727,31.79%的遗传变异存在于种群之间,68.21%的存在于种群内。本试验获得红三叶新品系的指纹图谱,可有效区分2份国内已登记红三叶品种。红三叶新品系Nei多样性指数0.0665,Shannon信息指数为0.1065,均低于平均水平,种群多样性相对较低。遗传距离表明红三叶新品系亲缘最近的3份材料依次为来自俄罗斯的60号>西班牙的22号>俄罗斯的53号。
5、红三叶新品系分株繁殖适宜切块6-7个,最高增殖系数为6.38,分枝期整枝扦插效率较高,最高增殖系数为6.40。组织培养研究表明：红三叶新品系种子消毒的最佳方法为75%酒精预处理,0.1%HgCl2消毒10min；最适无菌苗培养基为不添加激素的1/4MS+1.5%蔗糖；子叶、下胚轴、叶柄、叶片及根段在MS+0.5mg·L-16-BA+1～2mg·L-12,4-D培养基上都极易诱导出愈伤组织；最佳继代培养基为MS+0.5mg.L-16-BA+1.0mg·L-12,4-D;最佳分化培养基为MS+0.75mg·L-16-BA+0.5mg·L-1NAA,最高分化率为16.67%；茎芽增殖最佳培养基为MS+0.5mg·L-16-BA+1.0mg·L-1IBA;最佳生根条件为50mg·L-1浓度的IBA中浸泡1h后植入含0.5mg·L-1IBA的MS中,生根率达80%。1mg·L-1BA+0.5mg·L-1NAA激素配比适合红三叶茎尖和腋芽的萌发。
6、综合分析,初步推断内蒙古牙克石自然分布的红三叶可能起源于俄罗斯西部,属于欧洲血统。
The new line of red clover (Trifolium pratense L.) is an excellent cold resistant material suitable for alpine region in China, which filled the blank of red clover breeding in Inner Mongolia. Red clover new line and some red clover materials were studied by morphological features, chromosome, SSR markers, and asexual reproduction. The genetic origin of red clover which is natural distributed in Yakeshi in Inner Mongolia was discussed at first time in this paper. There is a great significance of the use and innovation of elite germplasm in red clover. Obtained the following results:
1、70materials can be divided into the early flowering, the middle flowering and late flowering, respectively, accounted for22.86%,68.57%and8.57%. According to growth period in second year, red clover can be divided into the early, medium and late maturity groups, and growth period were64d,71.96d and84.17d, respectively. New lines of red clover is a late flowering type. Growth period was74d belongs to the medium. Average growth rates in daily was0.44cm/d which close to the average level of red clover groups. Winter survival rates were from0%to88.89%in70materials at Hohhot. The new line of red clover winter rate was88.89%, and cold resistance was outstanding.
2、The studies of morphological characteristics have shown that red clover organs have great genetic variation. The coefficient of variation on leaf mark, branches, inflorescences per stem, seed color, internode length, lateral branch number, leaf area, natural height were relatively larger. The order of genetic diversity from the largest to the smallest was branches> leaves> seeds> flowers> roots. The crown width, height and branches were main indicators to divide morphological types of red clover. New lines of red clover had tan roots, and usually green stem which was semi-erect. Strains clump size was the middle in the red clover groups. Leaf mark were middle or no mark mainly. Leaf area was bigger relatively, and leaf shape was wider and shorter. The main flower color was purple,then light purple and pink, white occasionally. Seed color was relatively shallow, ana a large number of seeds was yellow. Seed size was middle, and1000-grain weight was smaller relative to the red clover groups. The new line of red clover was similar to the number of29,35,3,41,5,19,2,67in morphological characteristics.
3、Chromosome number of43red clover was2n-2x=14, x=7. Chromosome type included m, sm, st, and t, six of which had satellite. There were5kinds karyotype (1A,2A,1B,2B and3B)and8kinds of karyotype formula. The karyotype formula of new line of red clover was2n=2x=14=8m+6sm(2SAT). The asymmetry of interchromosomal is greater than intrachromosomal, and was relatively high in the red clover populations. According to the the Langlet's chromosome morphology criteria for the classification, the new line of red clover chromosome type is the T-type, primitive relatively.
4、The analysis of SSR marker showed that using8pairs of primers in32materials obtain219polymorphic bands totally. The polymorphism percentage was100%, and the number of effective alleles was1.1636. The Nei'diversity index, shannon information index, total variance, variance between the materials, coefficient of genetic differentiation and gene flow were0.1229,0.2241,0.1218,0.0387,0.3179respectively.31.79%genetic variation of the red clover exists in external populations,68.21%was in internal populations. The fingerprints of the red clover new line was obtained, which can effectively distinguish from two registered varieties of red clover in China. The Nei diversity index of red clover new line was0.0665, and Shannon's information index was0.1065, lower than the average population diversity level. SSR genetic distance showed that the new line of red clover was similar to the number of60,22,53, which were from Russia, Spain and Russia respectively.
5、The best ramets per plant was6-7in division propagation, and the highest proliferation coefficient was6.38; The highest proliferation coefficient of cutting can up to6.40using entire branches in branching period.The study of tissue culture showed:the best method of seeds sterilization for red clover is the75% ethyl alcohol treatment followed by disinfection with0.1%HgCl2for10min.1/4MS medium with1.5%sucrose without hormones was most effective for seed culture. Cotyledons, hypocotyl, petiole, leaf and root callus were easy to be induced in medium of MS containing0.5mg·L'16-BA and1-2mg·L-12,4-D. The best callus subculture medium was MS containing0.5mg·L-116-BA and1.0mg·L-12,4-D. The best medium for differentiation was MS plus0.75mg·L-16-BA and0.5mg·L-1NAA, the differentiation rate can reach up to16.67%. Corm-bud could be multiplicated greatly in MS medium plus0.5mg·L-16-BA and1.0mg·L-1IBA. Regenerated plants were immersed in50mg·L-1IBA for1h, then vaccinated in MS medium plus IBA0.5mg·L-1,the rooting rate was80%.1mg·L-16-BA+0.5mg·L-1NAA hormone combination was suitable for germination of shoot tips and axillary buds in red clover.
6、We concluded that red clover that distributed at Yakeshi in Inner Mongolia is likely to come from western Russia and was a European descent.
引文
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