东京野茉莉种子休眠特性的研究
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摘要
以浙江省遂昌县东京野茉莉新鲜种子为材料,通过对种子透水性、种被机械阻力、离体胚培养、发芽抑制物质提取及种子层积过程中的生理生化变化的研究,并利用赤霉素(GA3)进行发芽前预处理,探讨种子休眠机理及破除方法。结果表明:1)离体胚并无休眠特性,胚外组织中也不存在发芽抑制物质。尽管种子透水性影响其发芽率,但胚乳和种壳的机械束缚是东京野茉莉种子休眠的主要原因之一。2)层积后再机械处理可解除种子休眠。休眠能被赤霉素和(或)低温层积处理解除,说明种子同时存在生理抑制,其休眠类型属于综合性休眠。3)贮藏物质在层积过程中的动员与种子萌发率的提高密切相关,种内可溶性糖含量(R=0.885)、POD活性(R=0.658)与休眠的解除呈显著正相关。4)层积30 d后1 200 mg/L GA3处理的种子发芽率最高,为54%。胚的外覆物控制东京野茉莉的种子休眠,休眠解除的过程伴随种子内代谢水平的提高。
The fresh Styrax tonkinensis seeds were collected from Suichang county, Zhejiang province. Through the investigations on the physiological and biochemical changes in seed permeability, seed coat mechanical resistance, in vitro embryo culture, extraction of inhibitors of germination and seed stratification, and pretreatment with gibberellin(GA) before germination, the mechanism and release methods of seed dormancy were discussed. The results show that 1) the isolated embryos of the mature seeds had no characteristic of dormancy, and the leaching solution of the seed shell and endosperm did not contain germination inhibitors. The main reason causing seed dormancy was the mechanical obstacle of the seed shell and endosperm, while the permeability of the seed coat, which was not the main reason causing seed dormancy, also had some influences to seed germination; 2) Mechanical treatment after stratification can relieve seed dormancy. The dormancy could be released by gibberellin and(or) cold stratification, which suggested that there was the physiological dormancy in S.tonkinensis seeds, and the dormancy type was a comprehensive one; 3) the seed reserve mobilization tied closely to the dormancy release during cold stratification. In the process of breaking seed dormancy, the soluble sugar contents and POD activities showed an increasing trend along with the increasing of seed percentage germination, and the correlation coefficient was 0.885 and 0.658 respectively, which of both reached the significant level(P < 0.01); 4)The seed germination rate of 1 200 mg/L GA treated GAP after 30 days of stratification was the highest, which was 54%. The findings suggest that covering structure outside the S.tonkinenesis embryo controlled the seed dormancy, and the dormancy release was related to the up-regulation level of seed metabolism.
The fresh Styrax tonkinensis seeds were collected from Suichang county, Zhejiang province. Through the investigations on the physiological and biochemical changes in seed permeability, seed coat mechanical resistance, in vitro embryo culture, extraction of inhibitors of germination and seed stratification, and pretreatment with gibberellin(GA) before germination, the mechanism and release methods of seed dormancy were discussed. The results show that 1) the isolated embryos of the mature seeds had no characteristic of dormancy, and the leaching solution of the seed shell and endosperm did not contain germination inhibitors. The main reason causing seed dormancy was the mechanical obstacle of the seed shell and endosperm, while the permeability of the seed coat, which was not the main reason causing seed dormancy, also had some influences to seed germination; 2) Mechanical treatment after stratification can relieve seed dormancy. The dormancy could be released by gibberellin and(or) cold stratification, which suggested that there was the physiological dormancy in S.tonkinensis seeds, and the dormancy type was a comprehensive one; 3) the seed reserve mobilization tied closely to the dormancy release during cold stratification. In the process of breaking seed dormancy, the soluble sugar contents and POD activities showed an increasing trend along with the increasing of seed percentage germination, and the correlation coefficient was 0.885 and 0.658 respectively, which of both reached the significant level(P < 0.01); 4)The seed germination rate of 1 200 mg/L GA treated GAP after 30 days of stratification was the highest, which was 54%. The findings suggest that covering structure outside the S.tonkinenesis embryo controlled the seed dormancy, and the dormancy release was related to the up-regulation level of seed metabolism.
引文
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[3]柳新红,雷祖培,陈征海,等.浙江安息香属一新记录种:越南安息香[J].浙江农林大学学报,2012,29(2):319-320.
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[5]杨桦,唐仕斌,岳军伟,等.优良生物质能源树种东京野茉莉研究进展[J].安徽农业科学,2012,40(36):17648-17649,17657.
[6]PHUONG L X,SHIDA S,SAITO Y.Effects of heat treatment on brittleness of Styrax tonkinensis wood[J].Journal of Wood Science,2007,53(3):181-186.
[7]骆昱春,杨桦,曾志光,等.东京野茉莉木材性质分析与利用[J].江西农业大学学报,2007,29(1):77-80.
[8]柳新红,李因刚,何小勇.白花树研究进展[J].浙江林业科技,2008,28(5):61-65.
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[10]刘光斌,黄长干,刘苑秋,等.东京野茉莉油的提取及其制备生物柴油的初步研究[J].江西农业大学学报,2007,29(4):685-689.
[11]杨桦,龙蔚,骆昱春,等.芦溪岭林场东京野茉莉人工林3~5年生林木的种子产量及种子油食用价值研究[J].西部林业科学,2010,39(1):49-52.
[12]彭欢,吴亚丽,张子晗,等.东京野茉莉种子营养成分分析[J].林业工程学报,2015,29(3):56-58.
[13]范兰礼.东京野茉莉育苗和造林试验[J].江西林业科技,2001(4):3-4,8.
[14]许晓岗,丁芳芳,李翔,等.东京野茉莉种子休眠机制及其破除方法初探[J].西北植物学报,2012,32(11):2270-2278.
[15]李合生.植物生理生化实验原理和技术[M].北京:高等教育出版社,2000.
[16]唐安军.地涌金莲种子形态生理休眠及激素的动态变化[J].植物生理学报,2014(4):419-425.
[17]卢惊鸿.湖南牡丹资源调查及‘香丹’种子萌发研究[D].长沙:中南林业科技大学,2016.
[18]PUGLIA G,GRIMALDI S,CARTA A,et al.Pericarp structure of Glebionis coronaria(L.)Cass.ex Spach(Asteraceae)cypselae controls water uptake during germination[J].Seed Science Research,2015,25(3):255-266.
[19]耿文娟,冯贝贝,梅轩,等.野生欧洲李种子萌发特性[J].经济林研究,2017,35(1):20-25.
[20]BASKIN J M,BASKIN C C.A classification system for seed dormancy[J].Seed Science Research,2004,14(1):1-16.
[21]FOLEY M E.Leafy spurge(Euphorbia esula)seed dormancy[J].Weed Science,2004,52(1):74-77.
[22]FELIPPE G M.Qualea grandiflora:the seed and its germination[J].Revista Brasileira de Botanica,1990,13(1):33-37.
[23]BASKIN C C,BASKIN J M.Seeds:ecology,biogeography,and,evolution of dormancy and germination[M].California:Academic Press,1998.
[24]DEBEAUJON I,KOORNNEEF M.Gibberellin requirement for Arabidopsis seed germination is determined both by testa characteristics and embryonic abscisic acid[J].Plant Physiology,2000,122(2):415-424.
[25]BLACK M.Liberating the radicle:a case for softening-up[J].Seed Science Research,1996,6(2):39-42.
[26]GROOT S P C,KARSSEN C M.Gibberellins regulate seed germination in tomato by endosperm weakening:a study with gibberellin-deficient mutants[J].Planta,1987,171(4):525-531.
[27]JACOBSEN J V,PRESSMAN E,PYLIOTIS N A.Gibberellininduced separation of cells in isolated endosperm of celery seed[J].Planta,1987,129(2):113-122.
[28]TOOROP P E,BEWLEY J D,HILHORST H W M.Endo-β-mannanase isoforms are present in the endosperm and embryo of tomato seeds,but are not essentially linked to the completion of germination[J].Planta,1996,200(2):153-158.
[29]FU X X,LIU H N,XU J,et al.Primary metabolite mobilization and hormonal regulation during seed dormancy release in Cornus japonica var.chinensis[J].Scandinavian Journal of Forestry Research,2014,29(6):542-551.
[30]赵晓光.打破山桃种子休眠方法的研究[J].种子,2005,24(5):62-66.
[31]袁雅琪,钟秋平,袁小军,等.不同处理对3个种源漆树种子萌发的影响[J].经济林研究,2017,35(3):63-69.
[32]张军保.色木槭种子低温层积过程中贮藏物质和内源激素变化的研究[D].哈尔滨:东北林业大学,2008.
[33]郭明军,杨晓玲,张培玉,等.山楂种子休眠与萌发生理研究Ⅴ.山楂种子POD活性及其同工酶的变化与休眠的关系[J].河北科技师范学院学报,1997,21(3):25-28.
[34]ZHANG W,FAN J,TAN Q,et al.The effects of exogenous hormones on rooting process and the activities of key enzymes of Malus hupehensis stem cuttings[J].Plos One,2017,12(2):e0172320.
[35]LEWIS N G,YAMAMOTO E.Lignin:occurrence,biogenesis and biodegradation[J].Annual Review of Plant Physiology&Plant Molecular Biology,1990,41(1):455-96.
[36]PASSARDI F,COSIO C,PENEL C,et al.Peroxidases have more functions than a Swiss army knife[J].Plant Cell Reports,2005,24(5):255-265.
[37]王宝成,徐君霞,李明月,等.枫杨树种子休眠与萌发特性研究[J].森林工程,2018,34(2):11-15.
[2]PINYOPUSARERK K.Styrax tonkinensis:Taxonomy,ecology,silviculture and uses[R].AClAR Technical Report,1994.
[3]柳新红,雷祖培,陈征海,等.浙江安息香属一新记录种:越南安息香[J].浙江农林大学学报,2012,29(2):319-320.
[4]吴克选,曾志光,周小平,等.东京野茉莉野外调查报告[J].江西林业科技,2002(2):25-27.
[5]杨桦,唐仕斌,岳军伟,等.优良生物质能源树种东京野茉莉研究进展[J].安徽农业科学,2012,40(36):17648-17649,17657.
[6]PHUONG L X,SHIDA S,SAITO Y.Effects of heat treatment on brittleness of Styrax tonkinensis wood[J].Journal of Wood Science,2007,53(3):181-186.
[7]骆昱春,杨桦,曾志光,等.东京野茉莉木材性质分析与利用[J].江西农业大学学报,2007,29(1):77-80.
[8]柳新红,李因刚,何小勇.白花树研究进展[J].浙江林业科技,2008,28(5):61-65.
[9]顾雁蕾.越南安息香种子成分分析及超声波提油研究[D].杭州:浙江大学,2011.
[10]刘光斌,黄长干,刘苑秋,等.东京野茉莉油的提取及其制备生物柴油的初步研究[J].江西农业大学学报,2007,29(4):685-689.
[11]杨桦,龙蔚,骆昱春,等.芦溪岭林场东京野茉莉人工林3~5年生林木的种子产量及种子油食用价值研究[J].西部林业科学,2010,39(1):49-52.
[12]彭欢,吴亚丽,张子晗,等.东京野茉莉种子营养成分分析[J].林业工程学报,2015,29(3):56-58.
[13]范兰礼.东京野茉莉育苗和造林试验[J].江西林业科技,2001(4):3-4,8.
[14]许晓岗,丁芳芳,李翔,等.东京野茉莉种子休眠机制及其破除方法初探[J].西北植物学报,2012,32(11):2270-2278.
[15]李合生.植物生理生化实验原理和技术[M].北京:高等教育出版社,2000.
[16]唐安军.地涌金莲种子形态生理休眠及激素的动态变化[J].植物生理学报,2014(4):419-425.
[17]卢惊鸿.湖南牡丹资源调查及‘香丹’种子萌发研究[D].长沙:中南林业科技大学,2016.
[18]PUGLIA G,GRIMALDI S,CARTA A,et al.Pericarp structure of Glebionis coronaria(L.)Cass.ex Spach(Asteraceae)cypselae controls water uptake during germination[J].Seed Science Research,2015,25(3):255-266.
[19]耿文娟,冯贝贝,梅轩,等.野生欧洲李种子萌发特性[J].经济林研究,2017,35(1):20-25.
[20]BASKIN J M,BASKIN C C.A classification system for seed dormancy[J].Seed Science Research,2004,14(1):1-16.
[21]FOLEY M E.Leafy spurge(Euphorbia esula)seed dormancy[J].Weed Science,2004,52(1):74-77.
[22]FELIPPE G M.Qualea grandiflora:the seed and its germination[J].Revista Brasileira de Botanica,1990,13(1):33-37.
[23]BASKIN C C,BASKIN J M.Seeds:ecology,biogeography,and,evolution of dormancy and germination[M].California:Academic Press,1998.
[24]DEBEAUJON I,KOORNNEEF M.Gibberellin requirement for Arabidopsis seed germination is determined both by testa characteristics and embryonic abscisic acid[J].Plant Physiology,2000,122(2):415-424.
[25]BLACK M.Liberating the radicle:a case for softening-up[J].Seed Science Research,1996,6(2):39-42.
[26]GROOT S P C,KARSSEN C M.Gibberellins regulate seed germination in tomato by endosperm weakening:a study with gibberellin-deficient mutants[J].Planta,1987,171(4):525-531.
[27]JACOBSEN J V,PRESSMAN E,PYLIOTIS N A.Gibberellininduced separation of cells in isolated endosperm of celery seed[J].Planta,1987,129(2):113-122.
[28]TOOROP P E,BEWLEY J D,HILHORST H W M.Endo-β-mannanase isoforms are present in the endosperm and embryo of tomato seeds,but are not essentially linked to the completion of germination[J].Planta,1996,200(2):153-158.
[29]FU X X,LIU H N,XU J,et al.Primary metabolite mobilization and hormonal regulation during seed dormancy release in Cornus japonica var.chinensis[J].Scandinavian Journal of Forestry Research,2014,29(6):542-551.
[30]赵晓光.打破山桃种子休眠方法的研究[J].种子,2005,24(5):62-66.
[31]袁雅琪,钟秋平,袁小军,等.不同处理对3个种源漆树种子萌发的影响[J].经济林研究,2017,35(3):63-69.
[32]张军保.色木槭种子低温层积过程中贮藏物质和内源激素变化的研究[D].哈尔滨:东北林业大学,2008.
[33]郭明军,杨晓玲,张培玉,等.山楂种子休眠与萌发生理研究Ⅴ.山楂种子POD活性及其同工酶的变化与休眠的关系[J].河北科技师范学院学报,1997,21(3):25-28.
[34]ZHANG W,FAN J,TAN Q,et al.The effects of exogenous hormones on rooting process and the activities of key enzymes of Malus hupehensis stem cuttings[J].Plos One,2017,12(2):e0172320.
[35]LEWIS N G,YAMAMOTO E.Lignin:occurrence,biogenesis and biodegradation[J].Annual Review of Plant Physiology&Plant Molecular Biology,1990,41(1):455-96.
[36]PASSARDI F,COSIO C,PENEL C,et al.Peroxidases have more functions than a Swiss army knife[J].Plant Cell Reports,2005,24(5):255-265.
[37]王宝成,徐君霞,李明月,等.枫杨树种子休眠与萌发特性研究[J].森林工程,2018,34(2):11-15.