杨树雄性不育品种花粉发育过程的细胞学观察
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摘要
【目的】了解杨树小孢子败育的时期及影响小孢子败育的因素,为杨树雄性不育新品种选育提供科学依据。【方法】以杨树雄性不育品种‘泗杨1号’为材料,在光学显微镜下观察花粉粒细胞发育过程;在扫描电子显微镜下进行败育花粉粒的形态观察,并与可育品种‘南林3412’杨进行比较分析。【结果】显微镜下观测发现在四分体时期以后,即单核小孢子期,雄性不育品种‘泗杨1号’大部分小孢子发生异常,开始表现出败育性状。在花粉粒成熟期小孢子开始发生解体,小孢子败育。同时,用扫描电子显微镜观察仅有的少量花粉粒发现其异常变形,不同于正常花粉粒形态。【结论】‘泗杨1号’小孢子败育从单核小孢子时期开始,在花粉粒成熟期小孢子发生解体,小孢子败育。影响‘泗杨1号’小孢子败育主要原因是绒毡层细胞降解的延迟,影响小孢子发育营养物质的供给,导致小孢子发生败育;除此之外,花药室内的Ca2+浓度、花药内壁乌氏体的分布也有可能影响‘泗杨1号’小孢子的发育过程。
【Objective】Understand the period of microspore abortion and the factors affecting microspore abortion in poplar varieties to provide the scientific basis for breeding new male sterile poplar varieties.【Method】This paper used the male sterile Poplulus×euramericana 'Siyang-1'aged 16 years as study material. In spring 2016,the flower branches were collected with water training at room temperature. Then,we regularly sampled and fixed the material to make paraffin sections, and observe the cellular development of pollen grains under an Olympus-bx51 optical microscope.Moreover,we observed the morphology of abortive pollen grains under a Quanta 200 scanning electron microscope and compared it with that of the male fertile variety'Nanlin 3412.'【Result】The microspore development of'Siyang-1'began to develop abnormality after the tetrad period. In the mononuclear microspore phase,only a few cells formed mononuclear microspores,which were in a highly vacuolated or disintegrated state. The volume of microspores was obviously increased,showing signs of abortion. At the mature stage of pollen grains,the microspores began to disintegrate and the microspores were aborted. Only a small number of cells developed into two-celled pollen grains bonded to the remaining tapetum,which accumulated and clung to the inner wall of the anthers. The anthers shrank and dried,and no pollen was scattered. At the same time,the pollen grain morphology of'Siyang-1'was observed with the Quanta 200 scanning electron microscope. It was found that the pollen grains were abnormal and deformed,which was different from that of the normal pollen grains.【Conclusion】The abortion of'Siyang-1'began in the mononuclear microspore stage,and disintegrated at the mature stage of pollen grains,resulting in microspore abortion. The main reason affecting the abortion of'Siyang-1'was the delayed degradation of tapetum cells,which influenced the supply of nutrients to microspore development,resulting in microspore abortion. In addition,the concentration of Ca2+in the anther chamber and the distribu-tion of Uribe in the anther wall may also affect the process of microspore development. This paper provides a new idea for the study of male sterility of poplars,which is beneficial in the production practice in the future.
【Objective】Understand the period of microspore abortion and the factors affecting microspore abortion in poplar varieties to provide the scientific basis for breeding new male sterile poplar varieties.【Method】This paper used the male sterile Poplulus×euramericana 'Siyang-1'aged 16 years as study material. In spring 2016,the flower branches were collected with water training at room temperature. Then,we regularly sampled and fixed the material to make paraffin sections, and observe the cellular development of pollen grains under an Olympus-bx51 optical microscope.Moreover,we observed the morphology of abortive pollen grains under a Quanta 200 scanning electron microscope and compared it with that of the male fertile variety'Nanlin 3412.'【Result】The microspore development of'Siyang-1'began to develop abnormality after the tetrad period. In the mononuclear microspore phase,only a few cells formed mononuclear microspores,which were in a highly vacuolated or disintegrated state. The volume of microspores was obviously increased,showing signs of abortion. At the mature stage of pollen grains,the microspores began to disintegrate and the microspores were aborted. Only a small number of cells developed into two-celled pollen grains bonded to the remaining tapetum,which accumulated and clung to the inner wall of the anthers. The anthers shrank and dried,and no pollen was scattered. At the same time,the pollen grain morphology of'Siyang-1'was observed with the Quanta 200 scanning electron microscope. It was found that the pollen grains were abnormal and deformed,which was different from that of the normal pollen grains.【Conclusion】The abortion of'Siyang-1'began in the mononuclear microspore stage,and disintegrated at the mature stage of pollen grains,resulting in microspore abortion. The main reason affecting the abortion of'Siyang-1'was the delayed degradation of tapetum cells,which influenced the supply of nutrients to microspore development,resulting in microspore abortion. In addition,the concentration of Ca2+in the anther chamber and the distribu-tion of Uribe in the anther wall may also affect the process of microspore development. This paper provides a new idea for the study of male sterility of poplars,which is beneficial in the production practice in the future.
引文
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[4]张文超,曹媛,常童洁,等.毛白杨花粉不育过程中胼胝质的异常分布变化[J].东北林业大学学报,2013,41(1):68-71.DOI:10.3969/j.issn.1000-5382.2013.01.017.ZHANG W C,CAO Y,CHANG T J,et al. Abnormal distribution of corpus callosum during pollen sterility of Populus tomentosa[J]. Journal of Northeast Forestry University,2013,41(1):68-71.
[5]SNCHEZ D,VZQUEZ-SANTANA S. Embryology of Mammillaria dioica(Cactaceae)reveals a new male sterility phenotype[J]. Flora,2018,241:16-26. DOI:10.2985/026.019.0102.
[6]北大生物系植物遗传育种专业.雄性不育和雄性能育花药和花粉发育的细胞形态学观察[J].植物学报,1976,15(2):141-149.Biophyte Genetics and Breeding,Peking University. Cellular morphology of anther and pollen development in male sterility and male performance[J]. Journal of Plants, 1976, 15(2):141-149.
[7]彭婧,巩振辉,黄炜,等.辣椒雄性不育材料H9A小孢子败育机理[J].植物学报,2010(1):44-51.DOI:10.3969/j.issn.1674-3466.2010.01.006.PENG J,GONG Z H,HUANG W. et al. Mechanism of microspore sterility of pepper male sterile Line‘H9A’[J]. Journal of plants,2010,45(1):44-51.
[8]WANG A,XIA Q,XIE W,et al. The classical Ubisch bodies carry a sporophytically produced structural protein(RAFTIN)that is essential for pollen development[J]. Proceedings of the National Academy of Sciences,2003,100(24):14487-14492.DOI:10.1073/pnas.2231254100.
[9]石晶.乌氏体形成在花粉发育中的作用及研究进展[J].安徽农业科学,2009,37(33):16234-16263.DOI:10.13989/j.cnkj.0515-6611.2009.33.009.SHI J. Formation of Ubisch body and its function in the development of pollen and the related progress[J]. Journal of Anhui Agricultural Sciences,2009,37(33):16234-16263.
[10]李懿星,李莉,陈光辉.高等植物花药绒毡层发育研究进展[J].作物研究,2009,23(5):287-289.LI Y X,LI L,CHEN G H. Advances in the development of tapetal development in higher plant anther[J]. Crop Studies,2009,23(5):287-289.
[11] TIAN Y, XIAO S, LIU J, et al. MALE STERILE6021(MS6021)is required for the development of anther cuticle and pollen exine in maize[J]. Sci Rep,2017,7(1):16736. DOI:10.1038/s41598-017-16930-0.
[12]曾芳琴.油菜S45AB隐性核不育分子机理与应用研究[D].武汉:华中农业大学,2010.ZENG F Q. Studies on molecular mechanism and application of recessive genic male sterile rapeseed S45AB[D]. Wuhan:Huazhong Agricultural University,2010.
[13]林久生,王根轩.渗透胁迫诱导的小麦叶片细胞程序性死亡[J].植物生理与分子生物学学报,2001,27(3):221-225.LIN J S,WANG G X. Programmed cell death induced by osmotic stress in wheat leaves[J]. Journal of Plant Physiology and Molecular Biology,2001,27(3):221-225.
[14]HATANAKA M. Saturable and nonsaturable process of sugar uptake:effect of oncogenic transformation in transport and uptake of nutrients[J]. J Cell Physiol,1976,89(4):745-749. DOI:10.1002/jcp.1040890435.
[15]朱云,田惠桥.番茄花药发育过程中的多糖和脂滴的分布特征[J].园艺学报,2015,42(4):672-678. DOI:10. 16420/j.issn.0513-353x.2014-0886.ZHU Y,TIAN H Q. The distribution feature of polysaccharides and lipid droplets in the development of tomato anther[J]. Acta Horticalturae Sinica,2015,42(4):675-678.
[16]孟祥红,王建波,利容千.光周期对光敏胞质不育小麦花药发育过程中Ca2+分布的影响[J].植物学报,2000,42(1):446-454.MENG X H,WANG J B,LI R Q. Effect of photoperiocl on Ca2+distribution during anther delevlopment of cytoplasunic sterile wheat[J]. Acta Botanica Sinica,2000,42(1):446-454.
[2]樊汝汶,黄金生,李永敬.美洲黑杨无性系Ⅰ-63花粉粒的发育和超微结构[J].南京林产工业学院学报,1983(3):18-23+168-173.DOI:10.3969/j.issn.1000-2006.1983.08.003.FAN R W,HUANG J S,LI Y J. Pollen grain development and ultrastructure of Populus Americana cloneⅠ-63[J]. Journal of Nanjing Technologyica College of Forest Products,1983(3):18-23+168-173.
[3]康向阳.毛白杨花粉败育机制的研究[J].林业科学,2001,37(3):35-39.DOI:10.3321/j.issn.1001-7488.2001.03.007.KANG X Y. Studies on pollen abortion mechanism of Populus tomentosa[J]. Scientia Silvae Sinicae,2001,37(3):35-39.
[4]张文超,曹媛,常童洁,等.毛白杨花粉不育过程中胼胝质的异常分布变化[J].东北林业大学学报,2013,41(1):68-71.DOI:10.3969/j.issn.1000-5382.2013.01.017.ZHANG W C,CAO Y,CHANG T J,et al. Abnormal distribution of corpus callosum during pollen sterility of Populus tomentosa[J]. Journal of Northeast Forestry University,2013,41(1):68-71.
[5]SNCHEZ D,VZQUEZ-SANTANA S. Embryology of Mammillaria dioica(Cactaceae)reveals a new male sterility phenotype[J]. Flora,2018,241:16-26. DOI:10.2985/026.019.0102.
[6]北大生物系植物遗传育种专业.雄性不育和雄性能育花药和花粉发育的细胞形态学观察[J].植物学报,1976,15(2):141-149.Biophyte Genetics and Breeding,Peking University. Cellular morphology of anther and pollen development in male sterility and male performance[J]. Journal of Plants, 1976, 15(2):141-149.
[7]彭婧,巩振辉,黄炜,等.辣椒雄性不育材料H9A小孢子败育机理[J].植物学报,2010(1):44-51.DOI:10.3969/j.issn.1674-3466.2010.01.006.PENG J,GONG Z H,HUANG W. et al. Mechanism of microspore sterility of pepper male sterile Line‘H9A’[J]. Journal of plants,2010,45(1):44-51.
[8]WANG A,XIA Q,XIE W,et al. The classical Ubisch bodies carry a sporophytically produced structural protein(RAFTIN)that is essential for pollen development[J]. Proceedings of the National Academy of Sciences,2003,100(24):14487-14492.DOI:10.1073/pnas.2231254100.
[9]石晶.乌氏体形成在花粉发育中的作用及研究进展[J].安徽农业科学,2009,37(33):16234-16263.DOI:10.13989/j.cnkj.0515-6611.2009.33.009.SHI J. Formation of Ubisch body and its function in the development of pollen and the related progress[J]. Journal of Anhui Agricultural Sciences,2009,37(33):16234-16263.
[10]李懿星,李莉,陈光辉.高等植物花药绒毡层发育研究进展[J].作物研究,2009,23(5):287-289.LI Y X,LI L,CHEN G H. Advances in the development of tapetal development in higher plant anther[J]. Crop Studies,2009,23(5):287-289.
[11] TIAN Y, XIAO S, LIU J, et al. MALE STERILE6021(MS6021)is required for the development of anther cuticle and pollen exine in maize[J]. Sci Rep,2017,7(1):16736. DOI:10.1038/s41598-017-16930-0.
[12]曾芳琴.油菜S45AB隐性核不育分子机理与应用研究[D].武汉:华中农业大学,2010.ZENG F Q. Studies on molecular mechanism and application of recessive genic male sterile rapeseed S45AB[D]. Wuhan:Huazhong Agricultural University,2010.
[13]林久生,王根轩.渗透胁迫诱导的小麦叶片细胞程序性死亡[J].植物生理与分子生物学学报,2001,27(3):221-225.LIN J S,WANG G X. Programmed cell death induced by osmotic stress in wheat leaves[J]. Journal of Plant Physiology and Molecular Biology,2001,27(3):221-225.
[14]HATANAKA M. Saturable and nonsaturable process of sugar uptake:effect of oncogenic transformation in transport and uptake of nutrients[J]. J Cell Physiol,1976,89(4):745-749. DOI:10.1002/jcp.1040890435.
[15]朱云,田惠桥.番茄花药发育过程中的多糖和脂滴的分布特征[J].园艺学报,2015,42(4):672-678. DOI:10. 16420/j.issn.0513-353x.2014-0886.ZHU Y,TIAN H Q. The distribution feature of polysaccharides and lipid droplets in the development of tomato anther[J]. Acta Horticalturae Sinica,2015,42(4):675-678.
[16]孟祥红,王建波,利容千.光周期对光敏胞质不育小麦花药发育过程中Ca2+分布的影响[J].植物学报,2000,42(1):446-454.MENG X H,WANG J B,LI R Q. Effect of photoperiocl on Ca2+distribution during anther delevlopment of cytoplasunic sterile wheat[J]. Acta Botanica Sinica,2000,42(1):446-454.