二倍体芒小孢子发生及雄配子体发育研究
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摘要
该实验通过采集天然二倍体芒不同发育时期的幼穗,进行卡宝品红染色压片和石蜡切片制作,观察芒的花药发育过程,为芒的生殖生物学及其系统发育研究奠定理论基础。结果表明:(1)天然二倍体芒雄蕊3枚,雄蕊花药四室,花药壁由4层细胞组成,从外向内依次是表皮、药室内壁、中层、绒毡层,花药壁发育属于基本型;花药成熟时,中层和绒毡层降解消失,或者仅存痕迹,只剩表皮和纤维层细胞,但此时可观察到部分绒毡层降解延迟现象。(2)花粉母细胞减数分裂为连续型,成熟花粉粒为3-细胞型;花粉母细胞减数分裂后期Ⅱ出现染色体分裂不同步现象。(3)雄配子体发育过程中,同一花粉囊的花粉粒发育不同步;雌、雄配子体发育也不同步,且雄蕊先成熟。
The anther development process of natural diploid Miscanthus sinensis has been observed by carbol fuchsin staining and paraffin sectioning, after collecting young spikes of natural diploid M. sinensis at different stages. This experiment lay the theoretical foundation for reproductive biology and phylogeny of M. sinensis. The results as follows:(1) in natural diploid M. sinensis, there are 3 stamens and the anther is 4 sporangiates, which wall is composed of 4 layers: epidermis, endothecium, middle layer and tapetum. The development of anther wall belongs to the basic type. Middle layer and tapetum degenerated or just leavening the vestige, only epidermis and fibrous layer remained when maturity of the anther. However, there were some tapeyums degraded lingeringly at the time of anther matured.(2) The meiosis of pollen mother cells is of successive type and the mature pollen is 3-celled type. The phenomenon that chromosome asynchronous division in pollen mother cell meiosis anaphaseⅡ was observed.(3) The development of pollen grains of the same anther was not synchronous at the process of male gametogenesis. Male and female gametophytes also did not develop synchronously and stamens matured earlier than pistils.
The anther development process of natural diploid Miscanthus sinensis has been observed by carbol fuchsin staining and paraffin sectioning, after collecting young spikes of natural diploid M. sinensis at different stages. This experiment lay the theoretical foundation for reproductive biology and phylogeny of M. sinensis. The results as follows:(1) in natural diploid M. sinensis, there are 3 stamens and the anther is 4 sporangiates, which wall is composed of 4 layers: epidermis, endothecium, middle layer and tapetum. The development of anther wall belongs to the basic type. Middle layer and tapetum degenerated or just leavening the vestige, only epidermis and fibrous layer remained when maturity of the anther. However, there were some tapeyums degraded lingeringly at the time of anther matured.(2) The meiosis of pollen mother cells is of successive type and the mature pollen is 3-celled type. The phenomenon that chromosome asynchronous division in pollen mother cell meiosis anaphaseⅡ was observed.(3) The development of pollen grains of the same anther was not synchronous at the process of male gametogenesis. Male and female gametophytes also did not develop synchronously and stamens matured earlier than pistils.
引文
[1]中国科学院中国植物志编辑委员会.中国植物志[M].北京:科学出版社,1997,10:6-7.
[2]易自力.芒属能源植物资源的开发与利用[J].湖南农业大学学报(自然科学版),2012,38(5):455-463.YI Z L.Exploitation and utilization of Miscanthus as energy plant[J].Journal of Hunan Agricultural University(Natural Sciences),2012,38(5):455-463.
[3]刘明稀,蒋建雄,易自力.细胞工程技术在芒属能源作物上的应用[J].草业学报,2011,20(4):261-269.LIU M X,JIANG J X,YI Z L.Application of cell engineering in improving Miscanthus as a dedicated bioenergy crop[J].Acta Prataculture Sinica,2011,20(4):261-269.
[4]刁英,余作平,胡中立.芒属植物研究进展[J].现代农业科技,2011,(2):265-268.DIAO Y,YU Z P,HU Z L.Research advances on Miscanthus Anderss[J].Modern Agricultural Science and Technology,2011,(2):265-268.
[5]SANG T,ZHU W X.China’s bioenergy potential[J].Global Change Biology Bioenergy,2011,3(2):79-90.
[6]CHEN S L,RENVOIZE S A.Miscanthus[M]//WU Z Y,RAVEN P H,eds.Flora of China.Beijing&St.Louis:Science Press;Missouri Botanical Garden Press,2006:581-583.
[7]胡瑶,彭亮,赵盈盈,等.芒属植物在湖南长株地区园林绿化中的应用研究[J].湖南农业科学,2013,(15):102-104.HU Y,PENG L,ZHAO Y Y,et al.Application of Miscanthusin Changsha-Zhuzhou Regional landscape[J].Hunan Agricultural Sciences,2013,(15):102-104.
[8]任君霞,方佳,何勇清,等.芒属观赏草的研究进展[J].科技通报,2012,28(11):66-71.REN J X,FANG J,HE Y Q,et al.A review of research on ornamental grasses Miscanthus[J].Bulletin of Science and Technology,2012,28(11):66-71.
[9]KOBAYASHI K,YOKOI Y.Shoot population dynamics of persisting clones of Miscanthus sinensis in the warm-temperate region of Japan[J].Journal of Plant Research,2003,116(6):443-453.
[10]LAFFERTY J,LELLEY T.Cytogenetic studies of different Miscanthus species with potential for agricultural use[J].Plant Breeding,1994,113(3):246-249.
[11]GLOWACKA K,JRGENSEN U,KJELDSEN J B,et al.Can the exceptional chilling tolerance of C4 photosynthesis found in Miscanthus×giganteus be exceeded?Screening of a novel Miscanthus Japanese germplasm collection[J].Annals of Botany,2015,115(6):981-990.
[12]萧运峰,王锐,高洁.五节芒生态---生物学特性的研究[J].四川草原,1995,(1):25-29.XIAO Y F,WANG R,GAO J.Studies on ecological-biology of Miscanthus floridulus[J].Journal of Grassland and Forage Science,1995,(1):25-29.
[13]何立珍,贾大勇,洪亚辉,等.荻花粉母细胞减数分裂和花粉粒发育的研究[J].湖南农学院学报,1988,14(1):57-60.HE L Z,JIA D Y,HONG Y H,et al.Studies on the meiosis of pollen mother cells and the development of pollen grains of amur silvergrass[J].Journal of Hunan Agricultural College,1988,14(1):57-60.
[14]SOMKA A,KUTA E,PA Z·EK A,et al.Sterility of Miscanthus×giganteus results from hybrid incompatibility[J].Acta Biologica Cracoviensia Series Botanica,2012,54(1):1-8.
[15]邓果特.中国芒属植物染色体核型与倍性研究[D].长沙:湖南农业大学,2012.
[16]冯静仪,张振宇,李兴诗,等.一种优良的减数分裂实验材料及其制片[J].生物学通报,1986,(9):18.FENG J Y,ZHANG Z Y,LI X S,et al.An excellent meiotic material and its preparation[J].Bulletin of Biology,1986,(9):18.
[17]卢永根,冯九焕,刘向东,等.水稻(Oryza sativa L.)花粉及花药壁发育的超微结构研究[J].中国水稻科学,2002,16(1):29-37.LU Y G,FENG J H,LIU X D,et al.Ultrastructural studies on the developmental process of pollen and anther in rice(Oryza sativa L.)[J].Chinese Journal of Rice Science,2002,16(1):29-37.
[18]谭何新,文铁桥,张大兵.水稻花粉发育的分子机理[J].植物学通报,2007,24(3):330-339.TAN H X,WEN T Q,ZHANG D B.Molecular mechanisms of pollen development in Oryza sativa[J].Chinese Bulletin of Botany,2007,24(3):330-339.
[19]汤伟华,张蜀宁,孔艳娥.不结球白菜同源四倍体Pol CMS及其保持系花药发育的解剖学研究[J].西北植物学报,2008,28(4):704-708.TANG W H,ZHANG S N,KONG Y E.Anther development of Pol CMS and its corresponding maintainer line in autotetraploid non-heading Chinese cabbage[J].Acta Botanica Boreali-Occidentalia Sinica,2008,28(4):704-708.
[20]林树燕,丁雨龙.鹅毛竹大小孢子及雌雄配子体发育[J].西北植物学报,2012,32(5):907-914.LIN S Y,DING Y L.Development of the male and female gametophytes in Shibataea chinensis(Bambusoideae)[J].Acta Botanica Boreali-Occidentalia Sinica,2012,32(5):907-914.
[21]李秋莉,徐爱菊.高粱(Sorghum vulgare Pers.)雌、雄配子体形成过程的研究[J].辽宁师范大学学报(自然科学版),1997,20(2):66-69.LI Q L,XU A J.Study on the course of formation of male and female gametophyte in Sorghum vulgare Pers.[J].Journal of Liaoning Normal University(Natural Science),1997,20(2):66-69.
[22]卫星,申家恒.羊草大、小孢子发生与雌、雄配子体发育的观察[J].西北植物学报,2003,23(12):2 058-2 065.WEI X,SHEN J H.Observat ions on megasporogenesis,microsporogenesis and development of female and male gemetophyte of Aneurolepidium chinense(Trin.)Kitag[J].Acta Botanica Boreali-Occidentalia Sinica,2003,23(12):2 058-2 065.
[23]成仿云,金芝兰.谷子的大孢子发生和雌配子体发育[J].西北师范大学学报(自然科学版),1989,(2):58-63.CHENG F Y,JIN Z L.Megasporogenesis and development of female gametophyte in Setaria italica Beauv[J].Journol of Northwest Normal University(Natural Science),1989,(2):58-63.
[24]李进宇,张志翔,尹五元.连翘花的结构与繁育系统研究[J].西北植物学报,2006,26(8):1 548-1 553.LI J Y,ZHANG Z X,YIN W Y.Flower structure and reproduction system of Forsythia suspensa Vahl[J].Acta Botanica Boreali-Occidentalia Sinica,2006,26(8):1 548-1 553.
[25]赵元杰.芒繁育系统的研究[D].长沙:湖南农业大学,2010.
[2]易自力.芒属能源植物资源的开发与利用[J].湖南农业大学学报(自然科学版),2012,38(5):455-463.YI Z L.Exploitation and utilization of Miscanthus as energy plant[J].Journal of Hunan Agricultural University(Natural Sciences),2012,38(5):455-463.
[3]刘明稀,蒋建雄,易自力.细胞工程技术在芒属能源作物上的应用[J].草业学报,2011,20(4):261-269.LIU M X,JIANG J X,YI Z L.Application of cell engineering in improving Miscanthus as a dedicated bioenergy crop[J].Acta Prataculture Sinica,2011,20(4):261-269.
[4]刁英,余作平,胡中立.芒属植物研究进展[J].现代农业科技,2011,(2):265-268.DIAO Y,YU Z P,HU Z L.Research advances on Miscanthus Anderss[J].Modern Agricultural Science and Technology,2011,(2):265-268.
[5]SANG T,ZHU W X.China’s bioenergy potential[J].Global Change Biology Bioenergy,2011,3(2):79-90.
[6]CHEN S L,RENVOIZE S A.Miscanthus[M]//WU Z Y,RAVEN P H,eds.Flora of China.Beijing&St.Louis:Science Press;Missouri Botanical Garden Press,2006:581-583.
[7]胡瑶,彭亮,赵盈盈,等.芒属植物在湖南长株地区园林绿化中的应用研究[J].湖南农业科学,2013,(15):102-104.HU Y,PENG L,ZHAO Y Y,et al.Application of Miscanthusin Changsha-Zhuzhou Regional landscape[J].Hunan Agricultural Sciences,2013,(15):102-104.
[8]任君霞,方佳,何勇清,等.芒属观赏草的研究进展[J].科技通报,2012,28(11):66-71.REN J X,FANG J,HE Y Q,et al.A review of research on ornamental grasses Miscanthus[J].Bulletin of Science and Technology,2012,28(11):66-71.
[9]KOBAYASHI K,YOKOI Y.Shoot population dynamics of persisting clones of Miscanthus sinensis in the warm-temperate region of Japan[J].Journal of Plant Research,2003,116(6):443-453.
[10]LAFFERTY J,LELLEY T.Cytogenetic studies of different Miscanthus species with potential for agricultural use[J].Plant Breeding,1994,113(3):246-249.
[11]GLOWACKA K,JRGENSEN U,KJELDSEN J B,et al.Can the exceptional chilling tolerance of C4 photosynthesis found in Miscanthus×giganteus be exceeded?Screening of a novel Miscanthus Japanese germplasm collection[J].Annals of Botany,2015,115(6):981-990.
[12]萧运峰,王锐,高洁.五节芒生态---生物学特性的研究[J].四川草原,1995,(1):25-29.XIAO Y F,WANG R,GAO J.Studies on ecological-biology of Miscanthus floridulus[J].Journal of Grassland and Forage Science,1995,(1):25-29.
[13]何立珍,贾大勇,洪亚辉,等.荻花粉母细胞减数分裂和花粉粒发育的研究[J].湖南农学院学报,1988,14(1):57-60.HE L Z,JIA D Y,HONG Y H,et al.Studies on the meiosis of pollen mother cells and the development of pollen grains of amur silvergrass[J].Journal of Hunan Agricultural College,1988,14(1):57-60.
[14]SOMKA A,KUTA E,PA Z·EK A,et al.Sterility of Miscanthus×giganteus results from hybrid incompatibility[J].Acta Biologica Cracoviensia Series Botanica,2012,54(1):1-8.
[15]邓果特.中国芒属植物染色体核型与倍性研究[D].长沙:湖南农业大学,2012.
[16]冯静仪,张振宇,李兴诗,等.一种优良的减数分裂实验材料及其制片[J].生物学通报,1986,(9):18.FENG J Y,ZHANG Z Y,LI X S,et al.An excellent meiotic material and its preparation[J].Bulletin of Biology,1986,(9):18.
[17]卢永根,冯九焕,刘向东,等.水稻(Oryza sativa L.)花粉及花药壁发育的超微结构研究[J].中国水稻科学,2002,16(1):29-37.LU Y G,FENG J H,LIU X D,et al.Ultrastructural studies on the developmental process of pollen and anther in rice(Oryza sativa L.)[J].Chinese Journal of Rice Science,2002,16(1):29-37.
[18]谭何新,文铁桥,张大兵.水稻花粉发育的分子机理[J].植物学通报,2007,24(3):330-339.TAN H X,WEN T Q,ZHANG D B.Molecular mechanisms of pollen development in Oryza sativa[J].Chinese Bulletin of Botany,2007,24(3):330-339.
[19]汤伟华,张蜀宁,孔艳娥.不结球白菜同源四倍体Pol CMS及其保持系花药发育的解剖学研究[J].西北植物学报,2008,28(4):704-708.TANG W H,ZHANG S N,KONG Y E.Anther development of Pol CMS and its corresponding maintainer line in autotetraploid non-heading Chinese cabbage[J].Acta Botanica Boreali-Occidentalia Sinica,2008,28(4):704-708.
[20]林树燕,丁雨龙.鹅毛竹大小孢子及雌雄配子体发育[J].西北植物学报,2012,32(5):907-914.LIN S Y,DING Y L.Development of the male and female gametophytes in Shibataea chinensis(Bambusoideae)[J].Acta Botanica Boreali-Occidentalia Sinica,2012,32(5):907-914.
[21]李秋莉,徐爱菊.高粱(Sorghum vulgare Pers.)雌、雄配子体形成过程的研究[J].辽宁师范大学学报(自然科学版),1997,20(2):66-69.LI Q L,XU A J.Study on the course of formation of male and female gametophyte in Sorghum vulgare Pers.[J].Journal of Liaoning Normal University(Natural Science),1997,20(2):66-69.
[22]卫星,申家恒.羊草大、小孢子发生与雌、雄配子体发育的观察[J].西北植物学报,2003,23(12):2 058-2 065.WEI X,SHEN J H.Observat ions on megasporogenesis,microsporogenesis and development of female and male gemetophyte of Aneurolepidium chinense(Trin.)Kitag[J].Acta Botanica Boreali-Occidentalia Sinica,2003,23(12):2 058-2 065.
[23]成仿云,金芝兰.谷子的大孢子发生和雌配子体发育[J].西北师范大学学报(自然科学版),1989,(2):58-63.CHENG F Y,JIN Z L.Megasporogenesis and development of female gametophyte in Setaria italica Beauv[J].Journol of Northwest Normal University(Natural Science),1989,(2):58-63.
[24]李进宇,张志翔,尹五元.连翘花的结构与繁育系统研究[J].西北植物学报,2006,26(8):1 548-1 553.LI J Y,ZHANG Z X,YIN W Y.Flower structure and reproduction system of Forsythia suspensa Vahl[J].Acta Botanica Boreali-Occidentalia Sinica,2006,26(8):1 548-1 553.
[25]赵元杰.芒繁育系统的研究[D].长沙:湖南农业大学,2010.