Observation of the morphological and anatomical characteristics of male flower bud development in Litsea cubeba(Lour.) Pers.
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摘要: 采用体视显微镜、扫描电镜和石蜡切片技术对山鸡椒(Litsea cubeba(Lour.) Pers.)雄花花芽分化发育的外部形态和内部解剖结构进行了观察研究。结果显示:(1)山鸡椒雄花花芽分化发生可分为5个时期,即未分化期、花序原基分化期、苞片原基分化期、花原基分化期和花器官分化期,其中花器官分化期又可细分为花被原基分化期、雄蕊原基分化期和雌蕊原基分化期;各相邻分化时期存在一定重叠现象;花期从翌年1月上旬至3月下旬。(2)雄花成熟结构中具有独特的雄蕊蜜腺,蜜腺绿色且形态不规则,着生于内轮雄蕊基部,分布于花丝两侧,夹在内外轮雄蕊的花丝之间,与内轮花丝紧密相连。(3)雄蕊花药四室,花药壁发育属于基本型;腺质绒毡层;小孢子母细胞减数分裂过程中胞质分裂属于连续型;成熟花粉为2-细胞花粉粒;成熟花粉粒外壁刺突较多,刺突基部膨大,外壁露出部分粗糙,无薄壁区,有少数小穿孔。(4)山鸡椒雄花中绝大多数雌蕊发育至腹缝线卷合形成子房室时停止,柱头发育不良或者败育,花柱缩短或缺失,不能受精,直到开花结束,即发生退化。本研究明确了山鸡椒雄花花芽发育发生各个阶段时间、形态变化特点及外部形态变化特征,山鸡椒小孢子发生、雄配子体发育至散粉期变化特点和规律以及雄花中退化雌蕊发育的进程,可为山鸡椒优良品种选育、调控花期和提高结实率提供一定的参考。Abstract: To clarify the process of flower bud development in male flowers of Litsea cubeba(Lour.) Pers., we investigated the differentiation and development of the morphology and anatomy of male flower buds using stereomicroscopy, paraffin technology, and scanning electron microscopy. The results showed that: (1) The differentiation period of the male flower bud in L. cubeba was divided into five stages, including undifferentiation, inflorescence primordium differentiation, bract primordium differentiation, flower primordium differentiation, and floral organ differentiation, with the last stage further subdivided into perianth primordium differentiation, stamen primordium differentiation, and pistil primordium differentiation. There was some overlap in each differentiation stage. Flowering was from early January to late March the following year. (2) Mature L. cubeba males had unique irregular green stamen nectaries, which were borne at the base of the inner stamens, distributed on both sides of the filaments, intertwined with the filaments of the inner and outer stamens, and closely connected with the inner filaments. (3) Each anther had four pollen sacs and the development of the anther wall followed the basic model. The tapetum was glandular. Microspores were continuous during meiosis cytokinesis, and mature pollen grains were 2-pollen cells with very few microspore abortion during development. There were more spikes in ripe pollen exine, which was base dilated, and the exposed portion of the outer wall showed a rough, non-thin wall area, with a few small perforations. (4) Most pistils were arrested when the suture of abdominal cells formed into a peritoneal cavity. Therefore, the pistil had no stigma, and the style was shortened or missing,which process was called degradation. Fertilization did not happen until the end of flowering. The process was called degradation. We directly obtained the characteristics and patterns of male flower bud development, as well as the corresponding external morphogenesis, microsporogenesis, male gametophyte development, shedding, and development process of the pistillode in L. cubeba. These results provide a reference for the florescence regulation,selection and breeding of superior varieties of L. cubeba.
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Keywords:
- Litsea cubeba(Lour.) Pers. /
- Male flowers /
- Flower bud differentiation /
- Pistillode
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[1] Guo Q, Zeng KW, Gao XL, Zhu ZX, Zhang SY, Chai XY, Tu PF. Chemical constituents with NO production inhibitory and cytotoxic activities from Litsea cubeba[J]. J Nat Med, 2014, 69(1):94-99.
[2] 陈学恒. 我国山苍子资源利用现状和产业化前景评述[J]. 林业科学, 2003, 39(4):134-139. Chen XH. Comments on use of resources situation and industrial prospect of Litsea cubeba[J]. Scientia Silvae Sinicae, 2003, 39(4):134-139.
[3] 王旭, 杨关锋. 我国山苍子开发利用的现状与发展对策[J]. 经济林研究, 2010, 28(3):136-139. Wang X, Yang GF. Situation and development strategy of Litsea cubeba exploitation[J]. Nonwood Forest Research, 2010, 28(3):136-139.
[4] 崔永忠, 廖声熙, 崔凯, 刘方炎, 汪阳东, 陈益存. 贵州山区山苍子苗年生长规律[J]. 林业科学研究, 2013, 26(4):501-505. Cui YZ, Liao SX, Cui K, Liu FY, Wang YD, Chen YC. Study on the growth of one-year-old Litsea cubeba seedlings in mountainous area of Guizhou Province[J]. Forest Research, 2013, 26(4):501-505.
[5] 李芳, Yaru S, Howard H, Kevin O'S. 不同提取方法山苍子油的化学成分与抗氧化活性分析[J]. 林业科学, 2015, 51(3):124-131. Li F, Yaru S, Howard H, Kevin O'S. Chemical component and activity analysis of Litsea cubeba extracts obtained by different extraction methods[J]. Scientia Silvae Sinicae, 2015, 51(3):124-131.
[6] 曲波, 张薇, 陈旭辉, 李楠, 崔娜, 李天来. 植物花芽分化研究进展[J]. 中国农学通报, 2010, 26(24):109-114. Qu B, Zhang W, Chen XH, Li N, Cui N, Li TL. Plants flower bud differentiation research progress[J], Chinese Agricultural Science Bulletin, 2010, 26(24):109-114.
[7] 何燕红, 艾叶, 吴颖, 郭蕾, 包满珠. 孔雀草花芽分化和花药发育[J]. 华中农业大学学报, 2013, 32(2):18-24. He YH, Ai Y, Wu Y, Guo L, Bao MZ. Flower bud differentiation and anther development of Tagetes patula L.[J]. Journal of Huazhong Agricultural University, 2013, 32(2):18-24.
[8] 史洪琴. 大紫花芽形态分化观察[J]. 吉林农业:学术版, 2010(9):58-58. Shi HQ. Studies on morphological differentiation of big purple flower buds[J]. Jilin Agriculture:Academic Edition, 2010(9):58-58.
[9] 刘雄盛, 徐刚标, 梁文斌, 肖玉菲. 濒危植物水松小孢子发生和雄配子体发育的研究[J]. 植物科学学报, 2014, 32(1):58-66. Liu XS, Xu GB, Liang WB, Xiao YF. Studies microsporogenesis and development of male gametophytes in endangered species Glyptostrobus pensilis[J]. Plant Science Journal, 2014, 32(1):58-66.
[10] 许淑珺, 吴林芳, 胡晓颖, 徐信兰, 沈浩, 叶万辉. 桂味荔枝花器官的发生和发育过程研究[J]. 广西植物, 2012, 32(2):167-172. Xu SJ, Wu LF, Hu XY, Xu XL, Shen H, Ye WH. Floral organogenesis and development of Litchi chinensis Sonn. cv. Guiwei[J].Guihaia, 2012, 32(2):167-172.
[11] 陈英敏, 徐国辉, 王贺新, 朱玉, 王蜀, 刘家讯, 陈杉艳, 刘芳. 不同蓝莓品种花芽形成及二次开花特征[J]. 北方果树, 2016(4):6-10. Chen YM, Xu GL, Wang HX, Zhu Y, Wang S, Liu JS, Chen SB, Liu F. Flower bud formation and secondary flowering characteristics of different blueberry cultivars[J]. Northern Fruits, 2016(4):6-10.
[12] 高英, 董宁光, 张志宏, 张俊佩, 裴东. 早实核桃雌花芽分化外部形态与内部结构关系的研究[J]. 林业科学研究, 2010, 23(2):241-245. Gao Y, Dong NG, Zhang ZH, Zhang JP, Pei D. Relationship between external morphology and anatomical structure of precocious walnut during pistillate flower-bud diffe-rentiation[J].Forest Research, 2010, 23(2):241-245.
[13] 郝晨, 李云, 姜金仲, 刘洋. 四倍体刺槐大小孢子发育时期与花器形态的相关性[J]. 核农学报, 2006, 20(4):292-295. Hao C, Li Y, Jiang JZ, Liu Y. The correlation between micro and mega sporogenesis development and morphology of flower organ of tetraploid black locust[J]. Journal of Nuclear Agricultural Sciences, 2006, 20(4):292-295.
[14] 宋兴舜, 李发兵, 宋福南, 李开隆, 刘雪梅. 白桦雌花发育的形态相关[J]. 东北林业大学学报, 2008, 36(7):1-2. Song XS, Li FB, Song FN, Li KL, Liu XM. Morphological correlation of female flower development in Betula platyphylla[J]. Journal of Northeast Forestry University, 2008, 36(7):1-2.
[15] 胡正海. 植物分泌结构解剖学[M]. 上海:上海科学技术出版社, 2012:72-123. Hu ZH. Plant Secretory Structures Anatomy[M].Shanghai:Shanghai Science and Technology Press, 2012:72-123.
[16] 戴大临, 孔立生.山鸡椒雄花蜜腺形态发生的SEM研究[J].电子显微学报, 1993(1):98-98. Dai DL, Kong LS. Male nectaries morphogenesis SEM study of Litsea cubeba (Lour.) Pers[J].Journal of Chinese Electron Microscopy Society, 1993(1):98-98.
[17] Fahn A. Secretory Tissues in Plants[M]. London:Academic Press, 1979:51-113.
[18] Baum SF, Eshed Y, Bowman JL. The Arabidopsis nectary is an ABC-independent floralstructure[J]. Development, 2001, 128(22):4657-4667.
[19] 刘德胜, 方建民, 丁增发, 骆绪美. 山苍子的栽培和利用:上篇[J]. 安徽林业科技, 2011, 37(1):51-54. Liu DS,Fang JM, Ding ZF, Luo XM. Litsea cubeba cultivation and exploit:PartⅠ[J].Anhui Forestry Science and Technology, 2011, 37(1):51-54.
[20] 李捷, 李锡文. 世界樟科植物系统学研究进展[J]. 植物分类与资源学报, 2004, 26(1):1-11. Li J, Li XW. Advances in Lauraceae systematic research on the world scale[J]. Plant Diversity and Resources, 2004, 26(1):1-11.
[21] Bpm H. A revision of Lauraceae in Australia (excluding Cassytha)[J]. Aust Syst Bot, 1989, 2(2):135-367.
[22] Heo K, Werff HVD, Tobe H. Embryology and relationships of Lauraceae (Laurales)[J]. Bot J Linn Soc, 1998, 126(4):295-322.
[23] 胡适宜. 被子植物生殖生物学[M]. 北京:高等教育出版社, 2002:44-47. Hu SY. Angiosperms of Reproductive Biology[M]. Beijing:Higher Education Press, 2002:44-47. 24] 韩逸洋, 李利平, 陈发菊, 杨林森, 梁宏伟. 宜昌润楠大、小孢子发生及雌、雄配子体发育的研究[J]. 湖北农业科学, 2014(10):2352-2355. Han YY, Li LP, Chen FJ, Yang LS, Liang HW. The microsporogenesis, megasporogenesis, female and male gametophyte development of Machilus ichangensis Rehder & E. H. Wilson[J]. Hubei Agricultural Sciences, 2014(10):2352-2355.
[25] 周正立, 李志军, 龚卫江. 胡杨、灰叶胡杨开花生物学特性研究[J]. 武汉植物学研究, 2005, 23(2):163-168. Zhou ZL, LI ZJ, Gong WJ, Gao S. Study on the flowering phenology in Populus euphratica Olivier and Populus pruinosa Schrenk[J]. Journal of Wuhan Botanial Research, 2005, 23(2):163-168.
[26] 汤庚国, 向其柏. 樟科植物花粉形态研究[J]. 植物分类学报, 1995, 33(2):161-170. Tang GG, Xiang QB. Pollen morphology study of Laura-ceae[J]. Acta Phytotaxonomica Sinica, 1955, 33(2):161-170.
[27] Raj B, Werff HVD. A contribution to the pollen morphology of neotropical Lauraceae[J]. Ann Mo Bot Gard, 1988, 75(1):130-167.
[28] 孙坤, 陈家宽, 陈之端. 综述被子植物系统学中花发育研究的进展及对今后研究的思考[J]. 植物分类学报, 1998, 36(6):558-568. Sun K, Chen JK, Chen ZT. Progress in studies on floral development of angiosperms and some consideration on future studies[J]. Acta Phytotaxonomica Sinica,1998, 36(6):558-568.
[29] Wallander E, Albert VA. Phylogeny and classification of Oleaceae based on rps16 and trnL-F sequence data[J]. Am J Bot, 2000, 87(12):1827-1841.
[30] Wallander E. Systematics of Fraxinus L. (Oleaceae) and evolution of dioecy[J]. Plant Syst Evol, 2008, 273(1-2):25-49.
[31] Gleiser G, Verdu M. Repeated evolution of dioecy from androdioecy in Acer[J]. New Phytol, 2005, 165:633-640.
[32] Xu YC, Zhou LH, Hu SQ, Hao RM, Huang CJ, Zhao HB. The differentiation and development of pistils of hermaphrodites and pistillodes of males in androdioecious Osmanthus fragrans L. and implications for the evolution to androdioecy[J]. Plant Syst Evol, 2014, 300(5):843-849.
[33] Walkerlarsen J, Harder LD. The evolution of staminodes in angiosperms:patterns of stamen reduction, loss, and functional re-invention[J].Am J Bot, 2000, 87(10):1367-84.
[34] Hammerling GJ. Secondary pollen presentation in angiosperms and its biological significance[J]. Aust J Bot, 1993, 41(5):417-438.
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