为揭示板栗结实率低的内在原因,以河北迁西地区12年生 ‘燕山早丰’板栗为试材,采用石蜡切片、荧光显微观察和扫描电镜法,对其有性生殖发育过程进行了研究。结果表明:①4月下旬在1年生枝条的芽体内已具花原基,柔荑花序形成; 5月上旬完成性别分化; 5月中旬幼花药分化出表皮、内壁、中层及腺质绒毡层,花药壁发育为基本型; 5月下旬小孢子母细胞开始减数分裂,经过胞质同时分裂,形成四分体; 6月上中旬花药开裂,花粉粒散出,成熟花粉为二细胞、三沟。②雌花于5月上旬开始形成,分为雌花原基分化期、柱头原基分化期、柱头伸长期、子房形成期和开花期。6月中旬胚珠原基形成,子房8~9室,每室两个倒生胚珠,具双珠被,厚珠心。6月底至7月初胚囊成熟,属蓼型胚囊。③6月上中旬雄花开放后,花粉管经过约2周穿过花柱基部,于6月底至7月初开始进入胚囊,精卵融合为有丝分裂前型; 胚乳为核型胚乳; 合子经原胚棒状胚球形胚鱼雷型胚子叶胚,胚胎发育方式为柳叶菜型。8月上旬发育为成熟胚,8月底至9月初种子成熟时胚乳消失。④在雌配子发育过程中,7月中旬出现71.7%败育的胚珠,表现为空腔的胚囊、珠心组织退化等,这可能是‘燕山早丰’结实率低的原因之一。因此,阻止胚珠败育是提高板栗产量的关键。
Abstract
To investigate the causes of lack of fruit set in Chinese chestnut, a cultivar named ‘Yanshanzaofeng’ was employed in Qianxi County, Hebei province, and the process of sexual reproduction was examined by microscope. The results showed that ①catkins with floral primordia were formed in the buds of one-year olds shoots in later April. The sex differentiation was completed in early May. Later, the anther wall layers developed completely by mid-May and consisted of epidermis, endothecium, middle layers and glandular tapetum, and the anther wall was of the basic type. Microspore mother cells underwent meiosis through simultaneous cytokinesis in later May and gave rise to tetrads of microspores, which were tetrahedrally arranged. Mature pollens contained two cells with three germ pores. Anthers were dehiscent and pollen grains shed between early June and middle June. ② The female flowers were formed in early-May, and can be divided into female flower cluster primordium differentiation stage, alabastrum primordium differentiation stage, stigma primordium differentiation stage, stigma elongation stage, ovary formation stage and blooming stage. The ovule primordium was formed in middle June. The ovary has eight or nine locules, and each locule contained two anatropous ovules which were bitegmic and crassinucellate. The embryo sac was mature between later June and early July, and the development of embryo sac conformed to the Polygonum type. ③ When the anthers were dehiscent, the pollen tube passed through the basal style about two weeks. The pollen tube came into the embryo sac between later June and early July, and the syngamy was of premitotic type. The development of the endosperm was nuclear endosperm. The zygote developed from the proembryo to the clavate, globular, torpedo-shaped and cotyledon stage embryoid, and the development of embryo was of Onagrad type. ④The mature embryo formed in early August and the endosperm disappeared between later August and early September. The 71.7% abortive ovules were observed in the development of female gametophytes in middle July which showed the cavity of embryo and degeneration of nucellus tissue. The abortive ovules were identified as one of the major factors causing the low seed set in ‘Yanshanzaofeng’. Thus, It was the key to prevent the abortive ovules for the chestnut production.
{{custom_sec.title}}
{{custom_sec.title}}
{{custom_sec.content}}
参考文献
[1] 张宇和,刘銮,梁维坚,等.中国果树志.板栗 榛子卷[M].北京:中国林业出版社, 2005.
[2] Bounous G, Marinmi D T. Chestnut: botany, horticulture, and utilization[J]. Horticultural Reviews, 2005, 31: 291-347.
[3] 鄢丰霞,陈俊红.国内外的栗产业[J].落叶果树,2013,45(3):23-26.Yan F X, Chen J H. Domestic and abroad of chestnut industry [J]. Deciduous Fruits, 2013,45(3):23-26.
[4] 曹均,曹庆昌.板栗密植栽培实用技术[M].北京:中国农业科学技术出版社, 2010.
[5] Shi Z G, St?sser R. Reproductive biology of Chinese chestnut(Castanea mollissima Blume.)[J]. European Journal of Horticultural Science, 2005, 70(2): 96-113.
[6] Martín A, Herrera M A, Martín L M. In situ conservation and landscape genetics in forest species [J]. Journal of Natural Resources and Development, 2012(2): 1-5.
[7] 任立中,杨其光,杜国华.板栗花性别和器官分化的研究.Ⅱ:板栗生殖器官的发育和分化[J].安徽农学院学报,1981(2): 41-50.Ren L Z, Yang Q G, Du G H. Studies on the flower sex differentiation and organ differentiation of Chinese chestnut(Castanea mollissima BL.).II: The development and differentiation of reproductive organs of Chinese chestnut [J]. Journal of Anhui Agriculture College, 1981(2): 41-50.
[8] 张恒悦.板栗胚珠和胚的发育[J].山东农业大学学报,1986, 17(2):19-25.Zhang H Y. On the development of ovary and embryo of chestnut(Castanea mollissima BL.)[J]. Journal of Shandong Agricultural University, 1986, 17(2):19-25.
[9] 许慧玲,曹慧娟,李天庆.板栗(Castanea mollissima Bl.)的胚胎学研究(Ⅰ):胚珠、胚囊发育、受精和胚发生[J].北京林业大学学报,1988,10(1):10-16.Xu H L, Cao H J, Li T Q. Study on the embryologu of chestnut(Castanea mollissima BL.).I: The development of ovules and the embryo sac, fertilization and embryogenesis[J].Journal of Beijing Forestry University, 1988,10(1):10-16.
[10] 许慧玲,李天庆,曹慧娟.板栗(Castanea mollissima Bl.)的胚胎学研究(Ⅱ):雌配子体的超微结构[J].北京林业大学学报,1988, 10(3):91-96.Xu H L, Li T Q, Cao H J. Study on the embryologu of chestnut(Castanea mollissima BL.).II: The ultrastructure of the female gametophyte [J].Journal of Beijing Forestry University, 1988, 10(3):91-96.
[11] 姚家琳,夏仁学,马梦婷,等.板栗空苞形成因子的研究(Ⅱ):板栗空苞的胚胎学[J].华中农业大学学报,1989, 8(3):248-253.Yao J L, Xia R X, Ma M T et al. Study on factors of affecting empty-bur formation of Chinese chestnut(Castanea mollissima BL.)II. Embryology of empty-bur[J]. Journal of Huazhong Agricultural University, 1989, 8(3):248-253.
[12] 杜桂森,黄勤妮,李增彬,等.北京地区板栗胚珠发育的研究[J].北京师范学院学报:自然科学版,1992,13(3):51-54.Du G S, Huang Q N, Li Z B, et al. On the ovule growth of chestnut in Beijing area [J]. Journal of Beijing Teachers College, 1992,13(3):51-54.
[13] 夏立.中国板栗雌蕊形态结构与授粉特性的研究[D].昆明:西南林学院,2007.Xia L. Study on pistil morphology and pollination character of Chinese chestnut [D]. Kunming: Southwest Forestry College, 2007.
[14] 林翠翠.中国板栗花粉管生长及其机理研究[D].昆明:西南林学院,2008.Lin C C. Study on pollen tube growth and its mechanism of Chinese chestnut [D]. Kunming: Southwest Forestry College, 2008.
[15] Feng Y Q, Shen Y Y, Qin L et al. Short catkin 1, a novel mutant of Castanea mollissima, is associated with programmed cell death during chestnut staminate flower differentiation[J]. Scientia Horticulture, 2011, 130(2):431-435.
[16] Nakamura M. Fertilization and in vitro growth of ovules in Japanese Chestnut(Castanea crenata Sieb. et Zucc.)[J]. Journal of Japanese Society for Horticultural Science, 2003, 72(6):482-487.
[17] Nakamura M. Pollen tube growth and fertilization in Japanese chestnut(Castanea crenata Sieb. et Zucc.)[J]. Journal of Japanese Society for Horticultural Science, 2001, 70(5):561-566.
[18] Mert C, Soylu A. Flower and stamen structures of male-fertile and male-sterile chestnut(Castanea sativa Mill.)[J]. Journal of the American Society for Horticultural Science, 2006,131(6):752-759.
[19] Roberto Botta, Grazia Vergano, Giovanni Me, et al. Floralbiology and embryo development in chestnut(Castanea sativa Mill.)[J]. Hortscience, 1995, 30(6): 1283-1286.
[20] 樊莉丽,彭方仁,周琦,等.楸树大小孢子发生与雌雄配子体发育的研究[J].西北植物学报,2011,31(3):431-438.Fan L L, Peng F R, Zou Q, et al. Sporogensis and gametogensis of Catalpa bungei(Bignoniaceae)[J].Acta Botanica Boreali-Occidentalia Sinica, 2011,31(3):431-438.
[21] 郭素娟,吕文君,邹锋,等.迁西板栗主栽品种授粉组合的优化[J].江西农业大学学报,2013,35(3):437-443.Guo S J, Lyu W J, Zou F, et al. Experiment on screening pollination varieties for main Castanea mollissima varieties in Yan Mountain [J]. Acta Agriculturae Universitatis Jiangxiensis, 2013,35(3):437-443.
[22] 李和平.植物显微技术[M].2版.北京:科学出版社, 2009.
[23] 林树燕,郝娟娟,辛华,等.月月竹大、小孢子发生和雌、雄配子体发育研究[J].南京林业大学学报:自然科学版,2009,33(3):9-12.Lin S Y, Hao J J, Xin H, et al. The megasporgenesis, microsporgenesis and the development of their female and male gametophyte in Menstruocalamus sichuanesis [J]. Journal of Nangjing Forestry University: Natural Sciences Edition, 2009,33(3):9-12.
[24] 樊莉丽,彭方仁,王改萍,等.楸树自交及种内、种间杂交亲和性的细胞学观察[J].南京林业大学学报:自然科学版, 2013, 37(4): 1-7.Fan L L, Peng F R, Wang G P, et al. Cytological observation of fertilization compatibility of Catalpa bungei after self, intraspecific cross and interspecific cross-pollination [J]. Journal of Nangjing Forestry University:Natural Sciences Edition, 2013, 37(4):1-7.
[25] Mogensen H L. Ovule abortion in Quercus(Fagaceae)[J]. American Journal of Botany, 1975,62: 160-162.
[26]胡适宜.被子植物生殖生物学研究[M].北京:高等教育出版社,2005.
[27] 张林平,李保国,白志英,等.板栗雌花簇分化过程观察[J].果树科学,1999(4):280-283.Zhang L P, Li B G, Bai Z Y, et al. Observation of the female flower cluster differentiation process of chestnut [J]. Journal of Fruit Science, 1999(4):280-283.
[28] Deng M, Zhou Z K, Chen Y Q,et al. Systematic significance of the development and anatomy of flowers and fruit of Quercus schottkyana (Subgenus Cyclobalanopsis: Fagaceae)[J]. International Journal of Plant Sciences, 2008, 169(9): 1261-77.
[29] 郑诚乐,俞晓曲,潘东明,等.锥栗雌配子体形成与胚胎发育[J].福建农林大学学报:自然科学版,2009, 38(6):585-589.Zheng C L, Yu X Q, Pan D M, et al. Process of embryonic development and female gametophyte formation in Chinquapin [J].Journal of Fujian Agriculture and Forestry University:Natural Science Edition, 2009, 38(6):585-589.
[30] 白志英,路丙社,张林平,等.板栗雄花分化研究[J].经济林研究,2000,18(3):11-12.Bai Z Y, Liu B S, Zhang L P, et al. Study on the differentiation pattern of chestnut staminate flower[J]. Economic Forest Researches, 2000,18(3):11-12.
[31] 刘国彬,曹钧,周连第,等.板栗雄性不育的形态学观察[J].西南农业学报,2014,27(1):235-239.Liu G B, Cao J, Zhou L D, et al. Morphological observation on male sterility in Castanea mollissma Blume[J]. Southwest China Journal of Agricultural Sciences,2014,27(1):235-239.
[32] Omura M, Akihama T. Male sterility in chestnuts: A tentative plan for the seed propagation of fruit trees[J]. Gamma Field Symposia, 1980(19):77-89.
[33] Sisco P H, Hebard F V, Shi Y,et al. Cytoplasmic male sterility resulting from interspecific crosses in chestnut(Castanea spp.)[C]//Plant and Animal Genome Ⅸ. Conference, 2001.
[34] Kaul M L H. Male sterility in higher plants[M]. Berlin: Springer-Verlag, 1988.
[35] 熊彩珍,贾惠娟,何凤杰,等.槜李雌蕊形态及早期胚胎发育研究[J].果树学报,2010, 27(1):124-126.Xiong C Z, Jia H J, He F J, et al. Studies on the pistil morphology and early embryogenesie of Chinese plum cultivar Zuili [J]. Journal of Fruit Science, 2010, 27(1):124-126.
[36] Furukawa Y, Bukovac M J. Embryo sac development in Sour Cherry during the pollination period as related to fruit set[J]. Hort Science, 1989, 24(6):1005-1008.
[37] Burgos L, Egea J. Apricot embryo-sac development in relation to fruit set[J]. Journal of Horticultural Sciences, 1993, 68(2):203-208.
[38] Toyama T K. The pollen receptivity period and its relation to fruit setting in the stone fruits[J]. Fruit Varieties Journal, 1980,34(1):2-4.
[39] Reiser L, Fischer L R. The ovule and the embryo sac[J]. The Plant Cell, 1993,5(10):1291-1301.
[40] Jia H J, He F J, Xiong C Z, et al. Influences of cross pollination on pollen tube growth and fruit set in Zuili plums(Prunus salicina)[J]. Journal of Integrative Plant Biology, 2008,50(2):203-209.
[41] McKay J W. Self-sterility in the Chinese chestnut(Castanea mollissima)[J]. Proceedings of American Society for Horticultural Science, 1942,41: 156-161.
[42] Zheng C L, Pan D M, Zhang D C, et al. Studies on regularities of growth and development of Chinquapin(Castanea henryi)fruits[J].Acta Horticulturae, 2009(844):153.
基金
收稿日期:2014-01-08 修回日期:2015-03-09基金项目:“十二五”国家科技支撑计划(2013BAD14B0402); 高等学校博士学科点专项科研基金项目(20120014110011); 国家林业公益性行业科研专项重大项目(201204401) 第一作者:邹锋,博士生。*通信作者:郭素娟,教授。E-mail:gwangzs@263.net。引文格式:邹锋,郭素娟. 板栗‘燕山早丰’有性生殖过程解剖学研究[J]. 南京林业大学学报:自然科学版,2015,39(4):37-43.