极度濒危树种羊角槭花部形态特征及开花动态
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摘要
以天目山自然保护区人工栽培的濒危树种羊角槭(Acer yangjuechi Fang et P. L. Chiu)为研究对象,于花期观察了花部形态特征、单株与群体水平开花动态,分析开花与结实的关系,结果表明:1.羊角槭花杂性、顶生花序着生有雄花和两性花,两性花多位于花轴顶端,雄花位于侧生轴顶端或下部,花序雄花与两性花数量比为三角槭的2. 6倍,羊角槭枝条两性花偏少; 2.观察到两种形态雄花,Ⅰ型雄花子房微型、柱头极度短缩呈黑点状、花盘较平,Ⅱ型雄花子房退化成一团白色绒毛,2裂柱头隐现,花盘较厚、辐射状凹裂;两性花子房成扁球状突出,柱头显著伸出,花丝长度短于雄花; 3.单株两性花早于雄花开放,两性花集中于3日以内开放,呈集中开放模式,而对应此时同株上只有极个别雄花(7%以下)处于散粉阶段; 4.群体两性花开放时期略早于雄花,二者间存在相对的间隔期。由此对羊角槭开花与种胚发育影响因素进行了探讨。
To reveal the reproduction mechanism of low embryo development rate,the floral morphology and flowering process of Acer yangjuechi Fang et P. L. Chiu, the extremely endangered plants in Chinawas determined by testing trees cultivated in National Nature Reserve of Tianmu Mountain,Zhejiang Province. The results show that: (1) Bisexual flowers and maleflowers are inserted on a same apical inflorescence,and the bisexual flowers are always on the top of rachis while maleflowers on lateral rachis top or bottom. The quantity ratio of maleflowers to bisexual flowers is 6. 89 of 2. 6 times exceeding to A. buergerianum,which indicates less bisexual flowers of A. yangjuechi than A. buergerianum. (2) Two type of maleflowers were observed. Type Ⅰmaleflowers have tiny ovary,the stigma shortened extremely to a black spot and with flat flower disc while TypeⅡmaleflowers have biforked stigma outstanding upon ovary degenerated into white fluff and with thick and radial sunken flower disc. Bisexual flowers possess of outstanding oblate spheroid and shorter filament compared with maleflowers. (3) The flowering process of individual tree is of bisexual flowers flowering within 3 days previous to maleflowers while only under seven percent of maleflowers were on pollination stage at the same time. (4) For population flowering process,the flowering stage between bisexual flowers and maleflowers present an obvious interval and bisexual flowers flowering intensively previous to maleflowers. Therefore,the main cause of the high embryo abortion,which leads to endangered situation of A. yangjuechi,was discussed.
To reveal the reproduction mechanism of low embryo development rate,the floral morphology and flowering process of Acer yangjuechi Fang et P. L. Chiu, the extremely endangered plants in Chinawas determined by testing trees cultivated in National Nature Reserve of Tianmu Mountain,Zhejiang Province. The results show that: (1) Bisexual flowers and maleflowers are inserted on a same apical inflorescence,and the bisexual flowers are always on the top of rachis while maleflowers on lateral rachis top or bottom. The quantity ratio of maleflowers to bisexual flowers is 6. 89 of 2. 6 times exceeding to A. buergerianum,which indicates less bisexual flowers of A. yangjuechi than A. buergerianum. (2) Two type of maleflowers were observed. Type Ⅰmaleflowers have tiny ovary,the stigma shortened extremely to a black spot and with flat flower disc while TypeⅡmaleflowers have biforked stigma outstanding upon ovary degenerated into white fluff and with thick and radial sunken flower disc. Bisexual flowers possess of outstanding oblate spheroid and shorter filament compared with maleflowers. (3) The flowering process of individual tree is of bisexual flowers flowering within 3 days previous to maleflowers while only under seven percent of maleflowers were on pollination stage at the same time. (4) For population flowering process,the flowering stage between bisexual flowers and maleflowers present an obvious interval and bisexual flowers flowering intensively previous to maleflowers. Therefore,the main cause of the high embryo abortion,which leads to endangered situation of A. yangjuechi,was discussed.
引文
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3.方文培.中国植物志-第46卷槭树科[M].北京:科学出版社,1981:66-289.Fang W P.Flora of China Vol.46.Aceraceae[M].Beijing:Science Press,1981:66-289.
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5.钟泰林,李根有,石柏林.3种浙江特产濒危植物气体交换特征和叶绿素荧光特性研究[J].上海交通大学学报:农业科学版,2009,27(2):149-176.Zhong T L,Li G Y,Shi B L.Comparison of gas exchange and chlorophyll fluorescence parameters in three endangered species of Zhejiang province[J].Journal of Shanghai Jiaotong University:Agricultural Science,2009,27(2):149-176.
6.钱永生,王慧中,黎念林,等.十一种槭属植物遗传多样性AFLP分析[J].浙江林业科技,2007,27(1):1-5.Qian Y S,Wang H Z,Li N L,et al.Study on genetic diversity of 11 acer speciesby AFLPs[J].Journal of Zhejiang Forestry Science and Technology,2007,27(1):1-5.
7.李倩中,刘晓宏,苏家乐,等.槭属种质遗传多样性及亲缘关系的SRAP分析[J].江苏农业学报,2010,26(5):1032-1036.Li Q Z,Liu X H,Su J L,et al.Genetic diversity of Acer L.Germplasm revealed by sequence-related amplified polymorphism(SRAP)markers[J].Jiangsu Journal of Agricultural Sciences,2010,26(5):1032-1036.
8.陈香波,刘杨,赵明水,等.极度濒危树种羊角槭的种胚发育与休眠解除[J].林业科学,2017,53(4):65-73.Chen X B,Liu Y,Zhao M S,et al.Embryo development and dormancy releasing of Acer yangjuechi,the extremely endangered plant[J].Scientia Silvae Sinicae,2017,53(4):65-73.
9.Dafnia.Pollination ecology:a practical approach[M].New York:Oxford University Press,1992:1-57.
10.Dejong P C.Flowering and sex expression in Acer L.:abiosystematic study[R].Medelingen Nederland:Landbouwhogeschool Wageningen,1976.
11.Gleeson S K.Heterodichogamy in walnuts:inheritance and stable ratios[J].Evolution,1982,36(5):892-902.
12.Pendleton R L,Freeman D C,Mcarthur E D,et al.Gender specialization in heterodichogamous Grayia brandegei(Chenopodiaceae):evidence for an alternative pathway to dioecy[J].American Journal of Botany,2000,87(4):508-516.
13.Kimura M,Seiwa K,Suyama Y,et al.Flowering system of heterodichogamous Juglans ailanthifolia[J].Plant Species Biology,2003,18(2-3):75-84.
14.Bai W N,Zeng Y F,Zhang D Y.Mating patterns and pollen dispersal in a heterodichogamous tree,Juglans mandshurica(Juglandaceae)[J].New Phytologist,2007,176(3):699-707.
15.张灵芝,商辉,罗毅波,等.二重雌雄异型异熟物种色木槭3种表型花的细胞形态学研究[J].生物多样性,2011,19(5):551-557.Zhang L Z,Shang H,Luo Y B,et al.Morphology and cytology of three flower phenotypes in a duodichogamous tree species,Acer mono[J].Biodiversity Science,2011,19(5):551-557.
16.许小连,金荷仙,陈香波,等.濒危植物羊角槭小孢子发生与雄配子体发育研究[J].植物分类与资源学报,2012,34(4):339-346.Xu X L,Jin H X,Chen X B,et al.Studies on the formation of microspores and development of male gametophyte in A-cer yangjuechi(Aceraceae)[J].Plant Diversity and Resources,2012,34(4):339-346.
17.胡青,李凤兰,杜艺连,等.平基槭花性别分化的细胞形态学观察[J].植物研究,2009,29(2):136-140.Hu Q,Li F L,Du Y L,et al.Morphological and cytological study on floral sex differentiation of Acer truncatum Bge[J].Bulletin of Botanical Research,2009,29(2):136-140.
18.Hsu T Z,Chen Y S,Piet C,et al.Flora of China.Aceaceae[M].Beijing,China:Science Press,2008.
19.林立,林乐静,祝志勇,等.中国槭属植物的分类和系统发育研究[J].园艺学报,2017,44(8):1535-1547.Lin L,Lin L J,Zhu Z Y,et al.Studies on the taxonomy and molecular phylogeny of Acer in China[J].Acta Horticulturae Sinica,2017,44(8):1535-1547.
20.曹小勇,陈恺.濒危植物庙台槭生殖生物学研究1.花序和花的形态及发育[J].西北植物学报,1997,17(5):107-111.Cao X Y,Chen K.Studies on the reproductive biology of a China rare plant-Acer Miaotaiense1.Morphology and development of inflorescence and flower[J].Acta Botanica Boreali-OccidentaliaSinica,1997,17(5):107-111.
21.许小连.濒危植物羊角槭生殖生物学的初步研究[D].杭州:浙江农林大学,2012.Xu XL.Study on the reproductive biology of Acer yangjuechi[D].Hangzhou:Zhejiang A&FUniversity,2012.
2.潘跃芝,龚洵.濒危植物红花木莲大孢子发生和雌配子体发育的研究[J].西北植物学报,2002,22(5):1209-1214.Pan Y Z,Gong X.Megasporogenesis,development of female gametophyte of Manglietia insignis(Magnoliaceae)[J].Acta Botanica Boreali-Occidentalia Sinica,2002,22(5):1209-1214.
3.方文培.中国植物志-第46卷槭树科[M].北京:科学出版社,1981:66-289.Fang W P.Flora of China Vol.46.Aceraceae[M].Beijing:Science Press,1981:66-289.
4.裘宝林.浙江植物志(第4卷),冬青科-山茱萸科[M].杭州:浙江科学技术出版社,1993:56.Qiu B L.Flora of Zhejiang(vol.4):Aquifoliaceae-Cornaceae[M].Hangzhou:Zhejiang Science and Technology Press,1993:56.
5.钟泰林,李根有,石柏林.3种浙江特产濒危植物气体交换特征和叶绿素荧光特性研究[J].上海交通大学学报:农业科学版,2009,27(2):149-176.Zhong T L,Li G Y,Shi B L.Comparison of gas exchange and chlorophyll fluorescence parameters in three endangered species of Zhejiang province[J].Journal of Shanghai Jiaotong University:Agricultural Science,2009,27(2):149-176.
6.钱永生,王慧中,黎念林,等.十一种槭属植物遗传多样性AFLP分析[J].浙江林业科技,2007,27(1):1-5.Qian Y S,Wang H Z,Li N L,et al.Study on genetic diversity of 11 acer speciesby AFLPs[J].Journal of Zhejiang Forestry Science and Technology,2007,27(1):1-5.
7.李倩中,刘晓宏,苏家乐,等.槭属种质遗传多样性及亲缘关系的SRAP分析[J].江苏农业学报,2010,26(5):1032-1036.Li Q Z,Liu X H,Su J L,et al.Genetic diversity of Acer L.Germplasm revealed by sequence-related amplified polymorphism(SRAP)markers[J].Jiangsu Journal of Agricultural Sciences,2010,26(5):1032-1036.
8.陈香波,刘杨,赵明水,等.极度濒危树种羊角槭的种胚发育与休眠解除[J].林业科学,2017,53(4):65-73.Chen X B,Liu Y,Zhao M S,et al.Embryo development and dormancy releasing of Acer yangjuechi,the extremely endangered plant[J].Scientia Silvae Sinicae,2017,53(4):65-73.
9.Dafnia.Pollination ecology:a practical approach[M].New York:Oxford University Press,1992:1-57.
10.Dejong P C.Flowering and sex expression in Acer L.:abiosystematic study[R].Medelingen Nederland:Landbouwhogeschool Wageningen,1976.
11.Gleeson S K.Heterodichogamy in walnuts:inheritance and stable ratios[J].Evolution,1982,36(5):892-902.
12.Pendleton R L,Freeman D C,Mcarthur E D,et al.Gender specialization in heterodichogamous Grayia brandegei(Chenopodiaceae):evidence for an alternative pathway to dioecy[J].American Journal of Botany,2000,87(4):508-516.
13.Kimura M,Seiwa K,Suyama Y,et al.Flowering system of heterodichogamous Juglans ailanthifolia[J].Plant Species Biology,2003,18(2-3):75-84.
14.Bai W N,Zeng Y F,Zhang D Y.Mating patterns and pollen dispersal in a heterodichogamous tree,Juglans mandshurica(Juglandaceae)[J].New Phytologist,2007,176(3):699-707.
15.张灵芝,商辉,罗毅波,等.二重雌雄异型异熟物种色木槭3种表型花的细胞形态学研究[J].生物多样性,2011,19(5):551-557.Zhang L Z,Shang H,Luo Y B,et al.Morphology and cytology of three flower phenotypes in a duodichogamous tree species,Acer mono[J].Biodiversity Science,2011,19(5):551-557.
16.许小连,金荷仙,陈香波,等.濒危植物羊角槭小孢子发生与雄配子体发育研究[J].植物分类与资源学报,2012,34(4):339-346.Xu X L,Jin H X,Chen X B,et al.Studies on the formation of microspores and development of male gametophyte in A-cer yangjuechi(Aceraceae)[J].Plant Diversity and Resources,2012,34(4):339-346.
17.胡青,李凤兰,杜艺连,等.平基槭花性别分化的细胞形态学观察[J].植物研究,2009,29(2):136-140.Hu Q,Li F L,Du Y L,et al.Morphological and cytological study on floral sex differentiation of Acer truncatum Bge[J].Bulletin of Botanical Research,2009,29(2):136-140.
18.Hsu T Z,Chen Y S,Piet C,et al.Flora of China.Aceaceae[M].Beijing,China:Science Press,2008.
19.林立,林乐静,祝志勇,等.中国槭属植物的分类和系统发育研究[J].园艺学报,2017,44(8):1535-1547.Lin L,Lin L J,Zhu Z Y,et al.Studies on the taxonomy and molecular phylogeny of Acer in China[J].Acta Horticulturae Sinica,2017,44(8):1535-1547.
20.曹小勇,陈恺.濒危植物庙台槭生殖生物学研究1.花序和花的形态及发育[J].西北植物学报,1997,17(5):107-111.Cao X Y,Chen K.Studies on the reproductive biology of a China rare plant-Acer Miaotaiense1.Morphology and development of inflorescence and flower[J].Acta Botanica Boreali-OccidentaliaSinica,1997,17(5):107-111.
21.许小连.濒危植物羊角槭生殖生物学的初步研究[D].杭州:浙江农林大学,2012.Xu XL.Study on the reproductive biology of Acer yangjuechi[D].Hangzhou:Zhejiang A&FUniversity,2012.
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