杏花粉和花药壁发育的胚胎学特征
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摘要
为了解杏花药和花粉发育的主要特征及发育进程与花芽冬眠的关系,对不同发育阶段的花药和花粉进行了显微观察。结果表明,花药壁形成于11月上旬,由5层细胞构成,包括表皮、药室内壁、中层(含2层细胞)和绒毡层(分泌型)。小孢子母细胞形成于11月上旬,减数分裂完成于2月上旬,即小孢子发生过程历时约3个月。减数分裂开始时小孢子母细胞积累大量脂滴,减数分裂产生的4个小孢子在四分体内排列成四面体形。小孢子阶段约1个月,2–细胞花粉阶段约0.5个月,即雄配子体发育历时约1.5个月。3月初花蕾与芽鳞等长时小孢子发生有丝分裂产生2–细胞花粉。成熟花粉为2–细胞型,含脂滴而不含淀粉,属脂质花粉。小孢子母细胞形成时花芽开始冬眠,减数分裂完成时花芽结束冬眠,即小孢子发生与花芽冬眠同步。杏花粉发育历时4.5个月,其特征包括小孢子母细胞减数分裂时积累脂滴,小孢子发生与花芽冬眠同步,花蕾与芽鳞等长是雄配子体发生第一次有丝分裂的形态学标志。
In order to understand the embryological features of the pollen and anther wall of apricot,anthers of different ages were examined by means of light microscopy. The cell layers of anther wall were established in early November,including epidermis,endothecium,middle layers,and tapetum. The tapetum belonged to secretory type. The process of microsporogenesis lasted about 3 months. Microspore mother cells were formed in early November,and underwent meiosis during the period from late January to early February the next year. The microspore mother cell accumulated a large number of lipid droplets as meiosis began. The 4 meiotic nuclei were separated by simultaneous cytokinesis to form four microspores that were joined together in a tetrad. The four microspores in the tetrad were in a tetrahedral arrangement. The microspore was present for about one month before it underwent mitotic division to a 2-celled pollen grain. The 2-celled pollen stage lasted 2–3 weeks till anthesis. The mature pollen belonged to lipidic type,for it accumulated lipid droplets rather than starch. As the structure of anther walls was established,the scaly buds started winter dormancy,i.e.,they stopped elongation temporarily. When the meiosis of the microspore mother cell was completed,the winter dormancy was broken,i.e.,the scaly buds resumed elongation. Apparently,there was a coincidence between the mcirosporogenesis in the anther and the winter dormancy of the scaly buds. The microspore underwent mitotic division at the time when the flower bud reached the length of the scale. In summary,the process of apricot pollen development lasted 4.5 months,and the exclusive features included the coincidence between the microsporogenesis and the winter dormancy of the scaly buds,the accumulation of lipid droplets in the microspore mother cell,and that the first mitotic division of male gametophyte was marked by the growth of the flower bud to the length of the scale.
In order to understand the embryological features of the pollen and anther wall of apricot,anthers of different ages were examined by means of light microscopy. The cell layers of anther wall were established in early November,including epidermis,endothecium,middle layers,and tapetum. The tapetum belonged to secretory type. The process of microsporogenesis lasted about 3 months. Microspore mother cells were formed in early November,and underwent meiosis during the period from late January to early February the next year. The microspore mother cell accumulated a large number of lipid droplets as meiosis began. The 4 meiotic nuclei were separated by simultaneous cytokinesis to form four microspores that were joined together in a tetrad. The four microspores in the tetrad were in a tetrahedral arrangement. The microspore was present for about one month before it underwent mitotic division to a 2-celled pollen grain. The 2-celled pollen stage lasted 2–3 weeks till anthesis. The mature pollen belonged to lipidic type,for it accumulated lipid droplets rather than starch. As the structure of anther walls was established,the scaly buds started winter dormancy,i.e.,they stopped elongation temporarily. When the meiosis of the microspore mother cell was completed,the winter dormancy was broken,i.e.,the scaly buds resumed elongation. Apparently,there was a coincidence between the mcirosporogenesis in the anther and the winter dormancy of the scaly buds. The microspore underwent mitotic division at the time when the flower bud reached the length of the scale. In summary,the process of apricot pollen development lasted 4.5 months,and the exclusive features included the coincidence between the microsporogenesis and the winter dormancy of the scaly buds,the accumulation of lipid droplets in the microspore mother cell,and that the first mitotic division of male gametophyte was marked by the growth of the flower bud to the length of the scale.
引文
Bomblies K,Higgins J D,Yant L. 2015. Meiosis evolves:adaptation to external and internal environments. New Phytologist,208:306–323.
Borg M,Brownfield L,Twell D. 2009. Male gametophyte development:a molecular perspective. J Exp Bot,60:1465–1478.
Brewbaker J L.1967.The distribution and phylogenetic significance of binucleate and trinucleate pollen grains in the angiosperms.Am J Bot,54:1069-1083.
Chang F,Wang Y,Wang S,Ma H.2011.Molecular control of microsporogenesis in Arabidopsis.Curr Opin Plant Biol,14:66-73.
de Storme N,Copenhaver G P,Geelen D.2012.Production of diploid male gametes in Arabidopsis by cold-induced destabilization of postmeiotic radial microtubule arrays.Plant Physiol,160:1808-1826.
de Storme N,Geelen D.2014.The impact of environment stress on male reproductive development in plants:biological processes and molecular mechanisms.Plant Cell Environ,37:1-18.
Dickinson H G,Grant-Downton R.2009.Bridging the generation gap:flowering plant gametophytes and animal germlines reveal unexpected similarities.Biological Reviews,84:589-615.
Feng X Q,Dickinson H G.2007.Packaging the male germline in plants.Trends Genet,23:503-510.
Ge Shu-jun,Zhao Dan,Zhou Xue,Sun Qing,Liu Lin.2016.Cucumber leaf senescence causes in-situ and vacuolar degradation of lipid droplets.Acta Horticulturae Sinica,43(10):2021-2028.(in Chinese)葛姝君,赵丹,周雪,孙晴,刘林.2016.黄瓜叶片衰老过程中脂滴的原位降解和液泡内降解.园艺学报,43(10):2021-2028.
Gómez J F,Talle B,Wilson Z A.2015.Anther and pollen development:a conserved developmental pathway.J Integr Plant Biol,57:876-891.
Gong Yan-hui,Liu Lin.2015.Microchemical and ultramicroscopic observation on dynamics of storage reserves in chili pepper fruits during ripening.Acta Horticulturae Sinica,42(10):1944-1952.(in Chinese)公颜慧,刘林.2015.辣椒成熟过程中贮藏物质变化的显微化学与电镜观察.园艺学报,42(10):1944-1952.
Gurusamy C,Davis P J,Bal A K.2000.Seasonal changes in perennial nodules of beach pea(Lathyrus maratimus[L]Bigel.)with special reference to oleosomes.Int J Plant Sci,161:631-638.
Hong M.2005.Molecular genetic analyses of microsporogenesis and microgametogenesis in flowering plants.Annu Rev Plant Biol,56:393-434.
Hu Shi-yi.1982.Embryology of the angiosperms.Beijing:Higher Education Press:20-36.(in Chinese)胡适宜.1982.被子植物胚胎学.北京:高等教育出版社:20-36.
Jansson S,Douglas C J.2007.Populus:a model system for plant biology.Annu Rev Plant Biol,58:435-458.
Jones M L.2002.Lipids//Bancroft J D,Gamble M.Theory and practice of histological and techniques.London:Churchill Livingstone:564-565.
Julian C,Rodrigo J,Herrero M.2011.Stamen development and winter dormancy in apricot(Prunus armeniaca).Ann Bot,108:617-625.
Litwin J A.1985.Light microscopic histochemistry on plastic sections.Prog Histochem Cyto,16:1-84.
Liu L.2012.Ultrastructural study on dynamics of plastids and mitochondria during microgametogenesis in watermelon.Micron,43:412-417.
Liu L.2013.Ultrastructural study on dynamics of lipid bodies and plastids during ripening of chili pepper fruits.Micron,46:43-50.
Liu L.2015.Ultramicroscopic examination of mature massulae of Habenaria arinaria(Orchidaceae).Micron,74:1-7.
Liu L.2016.Ultramicroscopic characterization of mature pollen grains of Habenaria sagittifer.Horticultural Plant Journal,2(1):50-54.
Liu Lin,Bai Huaju.2003.Microscopic study on watermelon anther and pollen grains.China Cucurbits and Vegetables,16(5):13-14.(in Chinese)刘林,白华举.2003.西瓜花药与花粉显微结构研究.中国西瓜甜瓜,16(5):13-14.
Liu Lin,Yuan Dong-mei,Zhang Song-huan,Zheng Hui-min,Wang Ting-liang,Li Chun-qi.2007.Study on the ultrastructure of mature watermelon pollen grains.Journal of Fruit Science,24(3):385-388.(in Chinese)刘林,员冬梅,张松焕,郑慧敏,王庭梁,李春琦.2007.西瓜成熟花粉的超微结构.果树学报,24(3):385-388.
Lu Ting,Wang Yiguang,Luo Yibo,Zhang Chao,Fu Jianxin,Dong Bin,Hu Shaoqing,Zhao Hongbo.2017.Comparison on flower bud differentiation and development of different seasons in Osmanthus fragrans‘Sijigui’.Acta Horticulturae Sinica,44(6):1145-1156.(in Chinese)陆婷,王艺光,骆义波,张超,付建新,董彬,胡绍庆,赵宏波.2017.‘四季桂’不同季节的花芽分化与发育比较.园艺学报,44(6):1145-1156.
Murgia M,Charzynska M,Rougier M,Cresti M.1991.Secretory tapetum of Brassica oleracea L.:polarity and ultrastructural features.Sex Plant Reprod,4:28-35.
Murphy D J.2012.The dynamic roles of lipid bodies in animals,plants and microorganisms.Prog Lipid Res,40:43-50.
Pacini E,Bellani L M,Lozzi R.1986.Pollen,tapetum and anther development in two cultivars of sweet cherry(Prunus avium).Phytomorphology,36:197-210.
Pacini E,Juniper B E.1983.The ultrastructure of the formation and development of the amoeboid tapetum in Arum italicum Miller.Protoplasma,117:116-129.
Pacini E.1996.Types and meaning of pollen carbohydrate reserves.Sex Plant Reprod,9:362-366.
Pacini E.2010.Relationships between tapetum,loculus,and pollen during development.Int J Plant Sci,171:1-11.
Pyc M,Cai Y,Greer M S,Yurchenko O,Chapman K D,Dyer J M,Mullen R T.2017.Turning over a new leaf in lipid droplet biology.Trends Plant Sci,22:596-609.
Rodríguez-García M I,M'rani-Alaoui M,Fernández M C.2007.Behavior of storage lipids during development and germination of olive(Olea europaea L.)pollen.Protoplasma,221:237-244.
Scott R J,Spielman M,Dickinson H G.2004.Stamen structure and function.Plant Cell,16:S46-S60.
van der Schoot C,Rinne P L H.2011.Dormancy cycling at the shoot apical meristem:transitioning between self-organization and self-arrest.Plant Sci,180:120-131.
Wang Xin-chai.1986.Plant microtechnique.Fuzhou:Fujian Education Press:178-179.(in Chinese)王心钗.1986.植物显微技术.福州:福建教育出版社:178-179.
Wang Yong,Li Qiaoxia,Sun Kun,Chen Wen.2017.The study on dimorphic flower development and the soluble sugar and starch content in Viola philippica.Acta Horticulturae Sinica,44(2):323-329.(in Chinese)王镛,李巧峡,孙坤,陈纹.2017.紫花地丁开放花与闭锁花的发育及可溶性糖与淀粉含量的研究.园艺学报,44(2):323-329.
Wilson Z A,Zhang D B.2009.From Arabidopsis to rice:pathways in pollen development.J Exp Bot,60:1479-1492.
Winiarczyk K,Gebura J.2017.Formation of a unique structure during microsporogenesis in Tinantia anomala(Commelinaceae)anthers.Protoplasma,254:785-790.
Zhang Tao,Dong Wen-cai,Lin Lin.2013.Anatomical observation of pollen grain development in pumpkin.China Cucurbits and Vegetables,26(6):44-46.(in Chinese)张涛,董文才,刘林.2013.南瓜花粉发育解剖观察.中国瓜菜,26(6):44-46.
Zhu Yun,Tian Hui-qiao.2015.The distribution features of polysaccharide and lipid drops in the development of tomato anther.Acta Horticulturae Sinica,42(4):672-678.(in Chinese)朱云,田惠桥.2015.番茄花药发育中多糖和脂滴的分布特征.园艺学报,42(4):672-678.
Borg M,Brownfield L,Twell D. 2009. Male gametophyte development:a molecular perspective. J Exp Bot,60:1465–1478.
Brewbaker J L.1967.The distribution and phylogenetic significance of binucleate and trinucleate pollen grains in the angiosperms.Am J Bot,54:1069-1083.
Chang F,Wang Y,Wang S,Ma H.2011.Molecular control of microsporogenesis in Arabidopsis.Curr Opin Plant Biol,14:66-73.
de Storme N,Copenhaver G P,Geelen D.2012.Production of diploid male gametes in Arabidopsis by cold-induced destabilization of postmeiotic radial microtubule arrays.Plant Physiol,160:1808-1826.
de Storme N,Geelen D.2014.The impact of environment stress on male reproductive development in plants:biological processes and molecular mechanisms.Plant Cell Environ,37:1-18.
Dickinson H G,Grant-Downton R.2009.Bridging the generation gap:flowering plant gametophytes and animal germlines reveal unexpected similarities.Biological Reviews,84:589-615.
Feng X Q,Dickinson H G.2007.Packaging the male germline in plants.Trends Genet,23:503-510.
Ge Shu-jun,Zhao Dan,Zhou Xue,Sun Qing,Liu Lin.2016.Cucumber leaf senescence causes in-situ and vacuolar degradation of lipid droplets.Acta Horticulturae Sinica,43(10):2021-2028.(in Chinese)葛姝君,赵丹,周雪,孙晴,刘林.2016.黄瓜叶片衰老过程中脂滴的原位降解和液泡内降解.园艺学报,43(10):2021-2028.
Gómez J F,Talle B,Wilson Z A.2015.Anther and pollen development:a conserved developmental pathway.J Integr Plant Biol,57:876-891.
Gong Yan-hui,Liu Lin.2015.Microchemical and ultramicroscopic observation on dynamics of storage reserves in chili pepper fruits during ripening.Acta Horticulturae Sinica,42(10):1944-1952.(in Chinese)公颜慧,刘林.2015.辣椒成熟过程中贮藏物质变化的显微化学与电镜观察.园艺学报,42(10):1944-1952.
Gurusamy C,Davis P J,Bal A K.2000.Seasonal changes in perennial nodules of beach pea(Lathyrus maratimus[L]Bigel.)with special reference to oleosomes.Int J Plant Sci,161:631-638.
Hong M.2005.Molecular genetic analyses of microsporogenesis and microgametogenesis in flowering plants.Annu Rev Plant Biol,56:393-434.
Hu Shi-yi.1982.Embryology of the angiosperms.Beijing:Higher Education Press:20-36.(in Chinese)胡适宜.1982.被子植物胚胎学.北京:高等教育出版社:20-36.
Jansson S,Douglas C J.2007.Populus:a model system for plant biology.Annu Rev Plant Biol,58:435-458.
Jones M L.2002.Lipids//Bancroft J D,Gamble M.Theory and practice of histological and techniques.London:Churchill Livingstone:564-565.
Julian C,Rodrigo J,Herrero M.2011.Stamen development and winter dormancy in apricot(Prunus armeniaca).Ann Bot,108:617-625.
Litwin J A.1985.Light microscopic histochemistry on plastic sections.Prog Histochem Cyto,16:1-84.
Liu L.2012.Ultrastructural study on dynamics of plastids and mitochondria during microgametogenesis in watermelon.Micron,43:412-417.
Liu L.2013.Ultrastructural study on dynamics of lipid bodies and plastids during ripening of chili pepper fruits.Micron,46:43-50.
Liu L.2015.Ultramicroscopic examination of mature massulae of Habenaria arinaria(Orchidaceae).Micron,74:1-7.
Liu L.2016.Ultramicroscopic characterization of mature pollen grains of Habenaria sagittifer.Horticultural Plant Journal,2(1):50-54.
Liu Lin,Bai Huaju.2003.Microscopic study on watermelon anther and pollen grains.China Cucurbits and Vegetables,16(5):13-14.(in Chinese)刘林,白华举.2003.西瓜花药与花粉显微结构研究.中国西瓜甜瓜,16(5):13-14.
Liu Lin,Yuan Dong-mei,Zhang Song-huan,Zheng Hui-min,Wang Ting-liang,Li Chun-qi.2007.Study on the ultrastructure of mature watermelon pollen grains.Journal of Fruit Science,24(3):385-388.(in Chinese)刘林,员冬梅,张松焕,郑慧敏,王庭梁,李春琦.2007.西瓜成熟花粉的超微结构.果树学报,24(3):385-388.
Lu Ting,Wang Yiguang,Luo Yibo,Zhang Chao,Fu Jianxin,Dong Bin,Hu Shaoqing,Zhao Hongbo.2017.Comparison on flower bud differentiation and development of different seasons in Osmanthus fragrans‘Sijigui’.Acta Horticulturae Sinica,44(6):1145-1156.(in Chinese)陆婷,王艺光,骆义波,张超,付建新,董彬,胡绍庆,赵宏波.2017.‘四季桂’不同季节的花芽分化与发育比较.园艺学报,44(6):1145-1156.
Murgia M,Charzynska M,Rougier M,Cresti M.1991.Secretory tapetum of Brassica oleracea L.:polarity and ultrastructural features.Sex Plant Reprod,4:28-35.
Murphy D J.2012.The dynamic roles of lipid bodies in animals,plants and microorganisms.Prog Lipid Res,40:43-50.
Pacini E,Bellani L M,Lozzi R.1986.Pollen,tapetum and anther development in two cultivars of sweet cherry(Prunus avium).Phytomorphology,36:197-210.
Pacini E,Juniper B E.1983.The ultrastructure of the formation and development of the amoeboid tapetum in Arum italicum Miller.Protoplasma,117:116-129.
Pacini E.1996.Types and meaning of pollen carbohydrate reserves.Sex Plant Reprod,9:362-366.
Pacini E.2010.Relationships between tapetum,loculus,and pollen during development.Int J Plant Sci,171:1-11.
Pyc M,Cai Y,Greer M S,Yurchenko O,Chapman K D,Dyer J M,Mullen R T.2017.Turning over a new leaf in lipid droplet biology.Trends Plant Sci,22:596-609.
Rodríguez-García M I,M'rani-Alaoui M,Fernández M C.2007.Behavior of storage lipids during development and germination of olive(Olea europaea L.)pollen.Protoplasma,221:237-244.
Scott R J,Spielman M,Dickinson H G.2004.Stamen structure and function.Plant Cell,16:S46-S60.
van der Schoot C,Rinne P L H.2011.Dormancy cycling at the shoot apical meristem:transitioning between self-organization and self-arrest.Plant Sci,180:120-131.
Wang Xin-chai.1986.Plant microtechnique.Fuzhou:Fujian Education Press:178-179.(in Chinese)王心钗.1986.植物显微技术.福州:福建教育出版社:178-179.
Wang Yong,Li Qiaoxia,Sun Kun,Chen Wen.2017.The study on dimorphic flower development and the soluble sugar and starch content in Viola philippica.Acta Horticulturae Sinica,44(2):323-329.(in Chinese)王镛,李巧峡,孙坤,陈纹.2017.紫花地丁开放花与闭锁花的发育及可溶性糖与淀粉含量的研究.园艺学报,44(2):323-329.
Wilson Z A,Zhang D B.2009.From Arabidopsis to rice:pathways in pollen development.J Exp Bot,60:1479-1492.
Winiarczyk K,Gebura J.2017.Formation of a unique structure during microsporogenesis in Tinantia anomala(Commelinaceae)anthers.Protoplasma,254:785-790.
Zhang Tao,Dong Wen-cai,Lin Lin.2013.Anatomical observation of pollen grain development in pumpkin.China Cucurbits and Vegetables,26(6):44-46.(in Chinese)张涛,董文才,刘林.2013.南瓜花粉发育解剖观察.中国瓜菜,26(6):44-46.
Zhu Yun,Tian Hui-qiao.2015.The distribution features of polysaccharide and lipid drops in the development of tomato anther.Acta Horticulturae Sinica,42(4):672-678.(in Chinese)朱云,田惠桥.2015.番茄花药发育中多糖和脂滴的分布特征.园艺学报,42(4):672-678.