马蹄莲属(Zantedeschia)品种指纹图谱及生殖生物学研究
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摘要
马蹄莲属（Zantedeschia）为天南星科（Araceae）植物，全属共7种2亚种。按其习性分为两组：一组为常绿组（Sect.1. Evergreen）,另一组为落叶组（Sect.2. Deciduous）。常绿组品种耐寒性、抗病性均较强，常见的仅一种，即：白花马蹄莲（Z.aethiopica）；落叶组品种通常称为彩色马蹄莲，色彩艳丽，品种繁多，是当前国际流行主栽的品种，已注册的商业品种多达150多个，其中常用品种60多个，多数品种是由美国、新西兰和荷兰等国家的马蹄莲专业公司或科研机构培育和开发的。
本研究以马蹄莲属88份品种资源为研究材料，从形态学观测入手，应用ISSR-PCR分子标记、石蜡切片、花粉的生活力测定、品种间杂交等技术方法，开展了品种资源的形态学、DNA指纹图谱分析和生殖生物学研究，详细观察描述了马蹄莲主要的形态特征，找出了马蹄莲一致性、稳定性和特异性性状，制定出“马蹄莲属植物新品种DUS测试指南”，提出了基于分子标记结果的品种群分级标准，对马蹄莲属植物大小孢子发生和雌雄配子体发育过程进行了研究，在扫描电镜下观测了花粉显微形态特征，进而进行花粉生活力测定萌发试验，验证了花粉的活力和萌发特性以及品种杂交的生物学特性。主要结论如下：
1.马蹄莲属品种的叶型有披针形、卵形、三角形和戟形；花色有白色系、黄色系、橙色系、粉色系、红色系、紫色系和双色系等7种类型；佛焰苞从俯视角度分为锐角形、钝角形和圆形三种类型。
2.马蹄莲品种性状具有良好的一致性、稳定性和特异性，经形态学性状综合比较分析，筛选出来具有良好一致性特征的39个性状，经与UPOV测试指南比较，起草制定了《植物新品种特异性、一致性和稳定性测试指南马蹄莲属》，已获准向农业部报批（见附录2）。
3.经正交试验设计统计分析，提出了马蹄莲的ISSR-PCR优化反应体系，即25μl反应体系中，含有10×PCR buffer2.5μl，Mg~(2+)2.0mmol/l，dNTPs0.20mmol/l，Taq酶1.5U，引物0.25μmol/l，DNA模板1.0ng/μl。从96对引物中筛选出了15对适合进行ISSR－PCR研究，共扩增出690个位点，其中435个为多态性位点，平均每对引物扩增出46个位点和29个多态性位点，多态性比率为63.04%。
4.运用UPGMA聚类法对ISSR-PCR分子标记结果进行分析，在相似系数0.15处可将88份材料分为常绿马蹄莲和落叶马蹄莲二组；在相似系数0.29处可将87个落叶马蹄莲品种划分为盆栽品种亚群、切花品种亚群和盆花切花兼用品种亚群，每个亚群内根据花色可分为白花品种、黄花品种、橙花品种、红花品种及紫花品种等，与品种的表型性状相一致。
5.筛选出了6对可用于构建马蹄莲品种指纹图谱的核心引物，并绘制了指纹图谱模型。
6.研究发现彩色马蹄莲品种‘Majestic Red’胚珠倒生，双珠被，厚珠心，具珠被绒毡层。大孢子母细胞减数分裂形成的四分体呈直线排列或T型，合点端的大孢子发育为功能大孢子，其余3个大孢子退化，胚囊为单孢子发生的蓼型胚囊发育方式。每花通常有雄蕊2-3个，蝶形花粉囊，每侧由2个小孢子囊构成。花药壁由外到内依次为表皮、药室内壁、中层和绒毡层，绒毡层属变形绒毡层类型，小孢子形成时胞质分裂为修饰性同时型，小孢子排列成十字形四分体，成熟花粉为二胞花粉粒。小孢子在四分体时期，绒毡层细胞提前降解，致使释放的小孢子不能正常发育。
7.经扫描电镜观测，18个马蹄莲品种的花粉大小、形状较一致，为长球体形，畸形率为5.88%～43.59%，花粉极轴长为28.8～39.5μm，赤道轴长为21.9～32.3μm，各品种P/E值范围为1.14～1.43。花粉外壁光滑，纹饰不明显，无萌发沟。
8.在25℃恒温培养箱内暗培养条件下，6个马蹄莲品种花粉萌发率和花粉管长度均在24h内达到最高值，其中尤以9h内生长为主，筛选出附加10%蔗糖和0.01%硼酸的培养基为花粉活力培养检测的最适宜配方。
Genus Zantedeschia is a member of family Araceae. There are7species and2subspecies inthis genus. According to growth habit, they are divided into two sections. Section1is calledEvergreen, another section is named Deciduous. Evergreen section has only one popular species,Z. aethiopica,with the advantages of good cold-tolerance and disease-resistant. Cultivars inDeciduous section are called Calla Lily because they are rich in color and types. Calla Lilycultivars are the current international popular main products. There are more than60cultivarswith almost worldwide use now among more than150registered commercial cultivars. Mostcultivars in this genus were bred by the companies and research institutes from United States,New Zealand and Netherlands.
In this research88cultivars resources were used as materials. Based on morphologyobservation and technology methods such as, ISSR-PCR molecular mark, paraffin slice, andpollen micro-observation and cultivars breeding, the morphology DNA fingerprint andreproductive biology were studied. The important morphological characters were observed anddescribed in detail, the guidelines for the Distinctess, Uniformity and Stability (DUS) test ofGenus Zantedeschia were filled out based on the analysis of distinctive, uniform and stabilitycharacteristics. Based on ISSR-PCR molecular mark results, classing standard for4cultivarsgroup were advanced. Megaspore and microspore and the development of gametophytes wereanalyzed. Microscopic morphological characteristics of pollen were observed, the pollen viabilityand germination characteristics were verified. The main conclusions are as follows:
1. The leave blade type can be divided into lanceolate, ovate, triangular and hastate. Spathecolors have7types such as white, yellow, orange, pink, red, purple, and bicolor. Spathe viewedfrom top divided into three types: acute, obtuse and rounded.
2. All samples have a good consistency, stability and specificity. After the consistencyanalysis of morphological traits,39characters were carried out for DUS test and with goodconsistency compared with UPOV test guidelines. Guidelines for the conduct of tests forDistinctness, Uniformity and Stability Genus Zantedeschia were formulated. The standarddraft for approval was send up to Ministry of Agriculture.
3. Orthogonal experimental design was conducted to establish the reaction system.The total25μl reaction solution includes2.5μl10×PCR buffer，2.0mmol/l Mg~(2+),0.20mmol/l dNTPs，1.5UTaq enzyme,0.25μmol/l primer and1.0ng/μl DNA template.15primers were selected from96pairs for ISSR-PCR reaction. These primers amplified690loci.435loci among them werepolymorphic. Average per pair of primers amplified29polymorphic loci out of46loci. Thepolymorphism ratio was63.04%.
4. Using the UPGMA cluster based on molecular markers of ISSR-PCR results,88samplescould be divided into two groups. Zantedeschia aethiopica and Calla lily at similar coefficient of0.15. At similar coefficient of0.29,87Calla cultivars could be divided into3sub-grops, thepot-flower sub-group, cut-flower sub-group and pot-cut flower sub-group. In each sub-groupthere are white, yellow, orange, red and purple color types. This result was similar withmorphological classification.
5. Six core primer were selected to establish the fingerprint of Zantedeschia cultivars. Thefingerprint model of samples was drawn.
6. The megasporogensis, microsporogensis and the development of female and malegametophytes of Zantedeschia ‘Majestic Red’ were observed through traditional paraffin sections.It’s ovule is anatropous, unitegmic, and crassinucellar with integument tapeta. The megasporemother cell meiosis to form a linear megaspore tetrad. Among them, The chalazal end megasporedevelops into the embryo sac mother cell，which developed to Polygonum type embryo-sac，another3megaspores degenerated. Each flower had2-3stamens, with butterfly-shaped pollensacs，ach side with2small sporangial. The wall of anther was completely developed, whichconsisted of four layers: epidermis, endothecium, middle layers and tapetum. The tapetum isdeformation type. The cytokinesis of microspore mother cell during meiosis was ofsimultaneaous type. Microspore tetrads are mainly cross. Pollen grains had2-celled when shed.During the Microspore Tetrad， tapetum cells Degradated in advance，so the released smallspores couldn’t develop normally
7. According to scanning electron microscope observation, pollens among all samples aresimilar in size and shape, prolate shape. The length of polar axis and equatorial axis was for28.8～39.5μ m and21.9～32.3μ m.the P/E values range from1.14～1.43. Pollen extine wassmooth without aperture, pollen sculpture was not obvious.
8. At25℃in the dark condition, pollen germination ratio and pollen tube length grew to thehighest value within24hours, especially in early9h. The suitable medium for cultivation ofpollen viability testing was mixed with another10%sucrose and0.01%H3BO3.
本研究以马蹄莲属88份品种资源为研究材料，从形态学观测入手，应用ISSR-PCR分子标记、石蜡切片、花粉的生活力测定、品种间杂交等技术方法，开展了品种资源的形态学、DNA指纹图谱分析和生殖生物学研究，详细观察描述了马蹄莲主要的形态特征，找出了马蹄莲一致性、稳定性和特异性性状，制定出“马蹄莲属植物新品种DUS测试指南”，提出了基于分子标记结果的品种群分级标准，对马蹄莲属植物大小孢子发生和雌雄配子体发育过程进行了研究，在扫描电镜下观测了花粉显微形态特征，进而进行花粉生活力测定萌发试验，验证了花粉的活力和萌发特性以及品种杂交的生物学特性。主要结论如下：
1.马蹄莲属品种的叶型有披针形、卵形、三角形和戟形；花色有白色系、黄色系、橙色系、粉色系、红色系、紫色系和双色系等7种类型；佛焰苞从俯视角度分为锐角形、钝角形和圆形三种类型。
2.马蹄莲品种性状具有良好的一致性、稳定性和特异性，经形态学性状综合比较分析，筛选出来具有良好一致性特征的39个性状，经与UPOV测试指南比较，起草制定了《植物新品种特异性、一致性和稳定性测试指南马蹄莲属》，已获准向农业部报批（见附录2）。
3.经正交试验设计统计分析，提出了马蹄莲的ISSR-PCR优化反应体系，即25μl反应体系中，含有10×PCR buffer2.5μl，Mg~(2+)2.0mmol/l，dNTPs0.20mmol/l，Taq酶1.5U，引物0.25μmol/l，DNA模板1.0ng/μl。从96对引物中筛选出了15对适合进行ISSR－PCR研究，共扩增出690个位点，其中435个为多态性位点，平均每对引物扩增出46个位点和29个多态性位点，多态性比率为63.04%。
4.运用UPGMA聚类法对ISSR-PCR分子标记结果进行分析，在相似系数0.15处可将88份材料分为常绿马蹄莲和落叶马蹄莲二组；在相似系数0.29处可将87个落叶马蹄莲品种划分为盆栽品种亚群、切花品种亚群和盆花切花兼用品种亚群，每个亚群内根据花色可分为白花品种、黄花品种、橙花品种、红花品种及紫花品种等，与品种的表型性状相一致。
5.筛选出了6对可用于构建马蹄莲品种指纹图谱的核心引物，并绘制了指纹图谱模型。
6.研究发现彩色马蹄莲品种‘Majestic Red’胚珠倒生，双珠被，厚珠心，具珠被绒毡层。大孢子母细胞减数分裂形成的四分体呈直线排列或T型，合点端的大孢子发育为功能大孢子，其余3个大孢子退化，胚囊为单孢子发生的蓼型胚囊发育方式。每花通常有雄蕊2-3个，蝶形花粉囊，每侧由2个小孢子囊构成。花药壁由外到内依次为表皮、药室内壁、中层和绒毡层，绒毡层属变形绒毡层类型，小孢子形成时胞质分裂为修饰性同时型，小孢子排列成十字形四分体，成熟花粉为二胞花粉粒。小孢子在四分体时期，绒毡层细胞提前降解，致使释放的小孢子不能正常发育。
7.经扫描电镜观测，18个马蹄莲品种的花粉大小、形状较一致，为长球体形，畸形率为5.88%～43.59%，花粉极轴长为28.8～39.5μm，赤道轴长为21.9～32.3μm，各品种P/E值范围为1.14～1.43。花粉外壁光滑，纹饰不明显，无萌发沟。
8.在25℃恒温培养箱内暗培养条件下，6个马蹄莲品种花粉萌发率和花粉管长度均在24h内达到最高值，其中尤以9h内生长为主，筛选出附加10%蔗糖和0.01%硼酸的培养基为花粉活力培养检测的最适宜配方。
Genus Zantedeschia is a member of family Araceae. There are7species and2subspecies inthis genus. According to growth habit, they are divided into two sections. Section1is calledEvergreen, another section is named Deciduous. Evergreen section has only one popular species,Z. aethiopica,with the advantages of good cold-tolerance and disease-resistant. Cultivars inDeciduous section are called Calla Lily because they are rich in color and types. Calla Lilycultivars are the current international popular main products. There are more than60cultivarswith almost worldwide use now among more than150registered commercial cultivars. Mostcultivars in this genus were bred by the companies and research institutes from United States,New Zealand and Netherlands.
In this research88cultivars resources were used as materials. Based on morphologyobservation and technology methods such as, ISSR-PCR molecular mark, paraffin slice, andpollen micro-observation and cultivars breeding, the morphology DNA fingerprint andreproductive biology were studied. The important morphological characters were observed anddescribed in detail, the guidelines for the Distinctess, Uniformity and Stability (DUS) test ofGenus Zantedeschia were filled out based on the analysis of distinctive, uniform and stabilitycharacteristics. Based on ISSR-PCR molecular mark results, classing standard for4cultivarsgroup were advanced. Megaspore and microspore and the development of gametophytes wereanalyzed. Microscopic morphological characteristics of pollen were observed, the pollen viabilityand germination characteristics were verified. The main conclusions are as follows:
1. The leave blade type can be divided into lanceolate, ovate, triangular and hastate. Spathecolors have7types such as white, yellow, orange, pink, red, purple, and bicolor. Spathe viewedfrom top divided into three types: acute, obtuse and rounded.
2. All samples have a good consistency, stability and specificity. After the consistencyanalysis of morphological traits,39characters were carried out for DUS test and with goodconsistency compared with UPOV test guidelines. Guidelines for the conduct of tests forDistinctness, Uniformity and Stability Genus Zantedeschia were formulated. The standarddraft for approval was send up to Ministry of Agriculture.
3. Orthogonal experimental design was conducted to establish the reaction system.The total25μl reaction solution includes2.5μl10×PCR buffer，2.0mmol/l Mg~(2+),0.20mmol/l dNTPs，1.5UTaq enzyme,0.25μmol/l primer and1.0ng/μl DNA template.15primers were selected from96pairs for ISSR-PCR reaction. These primers amplified690loci.435loci among them werepolymorphic. Average per pair of primers amplified29polymorphic loci out of46loci. Thepolymorphism ratio was63.04%.
4. Using the UPGMA cluster based on molecular markers of ISSR-PCR results,88samplescould be divided into two groups. Zantedeschia aethiopica and Calla lily at similar coefficient of0.15. At similar coefficient of0.29,87Calla cultivars could be divided into3sub-grops, thepot-flower sub-group, cut-flower sub-group and pot-cut flower sub-group. In each sub-groupthere are white, yellow, orange, red and purple color types. This result was similar withmorphological classification.
5. Six core primer were selected to establish the fingerprint of Zantedeschia cultivars. Thefingerprint model of samples was drawn.
6. The megasporogensis, microsporogensis and the development of female and malegametophytes of Zantedeschia ‘Majestic Red’ were observed through traditional paraffin sections.It’s ovule is anatropous, unitegmic, and crassinucellar with integument tapeta. The megasporemother cell meiosis to form a linear megaspore tetrad. Among them, The chalazal end megasporedevelops into the embryo sac mother cell，which developed to Polygonum type embryo-sac，another3megaspores degenerated. Each flower had2-3stamens, with butterfly-shaped pollensacs，ach side with2small sporangial. The wall of anther was completely developed, whichconsisted of four layers: epidermis, endothecium, middle layers and tapetum. The tapetum isdeformation type. The cytokinesis of microspore mother cell during meiosis was ofsimultaneaous type. Microspore tetrads are mainly cross. Pollen grains had2-celled when shed.During the Microspore Tetrad， tapetum cells Degradated in advance，so the released smallspores couldn’t develop normally
7. According to scanning electron microscope observation, pollens among all samples aresimilar in size and shape, prolate shape. The length of polar axis and equatorial axis was for28.8～39.5μ m and21.9～32.3μ m.the P/E values range from1.14～1.43. Pollen extine wassmooth without aperture, pollen sculpture was not obvious.
8. At25℃in the dark condition, pollen germination ratio and pollen tube length grew to thehighest value within24hours, especially in early9h. The suitable medium for cultivation ofpollen viability testing was mixed with another10%sucrose and0.01%H3BO3.
引文
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