硫素形态对苗期菘蓝生长生理特性及次生代谢的影响
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摘要
为探讨不同硫素形态对菘蓝生长、生理及次生代谢的影响,设置CK(无硫)、Na_2SO_4、Na_2SO_3、NaHSO_3、Na_2S和Na_2S_2O_3 6个试验处理,通过盆栽试验研究不同硫素形态对菘蓝生长、光合生理、氮硫代谢、营养物质和靛蓝、靛玉红含量的影响。结果表明,Na_2SO_4处理最有利于菘蓝叶片鲜重和干重的积累,NaHSO_3处理次之,与CK相比其他处理组对其生长和生物量积累无显著促进作用。硫素处理后叶片的光合色素含量及光合效率均显著高于无硫处理,其中以NaHSO_3处理后净光合速率最高,Na_2S处理叶绿素含量最高。不同形态硫素处理对氮硫代谢的影响显著不同,其中以Na_2SO_3处理对氮代谢酶活性促进作用最显著,Na_2SO_4处理可显著提高硫代谢酶活性,而缺硫(CK)对氮硫代谢酶活性无显著的抑制作用。叶片中营养物质含量对不同硫素的响应存在差异,其中缺硫显著降低叶片中可溶性蛋白和游离氨基酸含量,可溶性糖和可溶性蛋白含量以NaHSO_3处理下最高,而游离氨基酸含量以Na_2SO_4处理最高。不同于生理响应,次生代谢物质靛蓝和靛玉红含量以Na_2S处理最高。综合分析不同硫素形态对菘蓝叶采收品质的影响,以Na_2SO_4处理菘蓝后其生长状态处于最佳水平,且叶内活性成分也相对较高。可为菘蓝栽培选用适宜的硫素形态提供理论参考。
To investigate the effect of S in different chemical forms on the growth, physiology and secondary metabolism of Isatis indigotica, an experiment with six treatments [CK(no added S), Na_2SO_4, Na_2SO_3, NaHSO_3, Na_2S and Na_2S_2O_3] was set up, and the changes in leaf fresh and dry weight, photosynthetic physiology, nitrogen and sulfur metabolism, nutritional components and content of indigo and indirubin studied, using a pot trial. It was found that the fresh and dry weight were greatest in the Na_2SO_4 treatment, with the NaHSO_3 treatment being second highest. Compared with CK, the other S treatments did not significantly promote biomass accumulation. The net photosynthetic rates and chlorophyll contents in sulfur treatments were higher than those observed in CK, and the highest observed net photosynthetic rate and chlorophyll contents occurred in the NaHSO_3 and Na_2S, respectively. Na_2S_3 and Na_2SO_4 increased the activities of nitrogen and sulfur metabolism enzymes respectively compared to other treatments, with the CK being no inhibiting obvious in their effect. The responses of nutritional components to sulfur forms differed. Soluble protein and soluble amino acid concentrations were decreased in CK plants compared to S-treated plants, with the highest soluble protein, free amino acid, and soluble sugar concentrations observed in the NaHSO_3, Na_2SO_4 and NaHSO_3 treatments, respectively. In contrast to the physiological responses, the levels of secondary metabolites indigo and indirubin were the highest in Na_2S treatment. Considering both growth stimulation and secondary metabolite accumulation, the Na_2SO_4 treatment is recommended as the most beneficial in the cultivation of I. indigotica seedlings.
To investigate the effect of S in different chemical forms on the growth, physiology and secondary metabolism of Isatis indigotica, an experiment with six treatments [CK(no added S), Na_2SO_4, Na_2SO_3, NaHSO_3, Na_2S and Na_2S_2O_3] was set up, and the changes in leaf fresh and dry weight, photosynthetic physiology, nitrogen and sulfur metabolism, nutritional components and content of indigo and indirubin studied, using a pot trial. It was found that the fresh and dry weight were greatest in the Na_2SO_4 treatment, with the NaHSO_3 treatment being second highest. Compared with CK, the other S treatments did not significantly promote biomass accumulation. The net photosynthetic rates and chlorophyll contents in sulfur treatments were higher than those observed in CK, and the highest observed net photosynthetic rate and chlorophyll contents occurred in the NaHSO_3 and Na_2S, respectively. Na_2S_3 and Na_2SO_4 increased the activities of nitrogen and sulfur metabolism enzymes respectively compared to other treatments, with the CK being no inhibiting obvious in their effect. The responses of nutritional components to sulfur forms differed. Soluble protein and soluble amino acid concentrations were decreased in CK plants compared to S-treated plants, with the highest soluble protein, free amino acid, and soluble sugar concentrations observed in the NaHSO_3, Na_2SO_4 and NaHSO_3 treatments, respectively. In contrast to the physiological responses, the levels of secondary metabolites indigo and indirubin were the highest in Na_2S treatment. Considering both growth stimulation and secondary metabolite accumulation, the Na_2SO_4 treatment is recommended as the most beneficial in the cultivation of I. indigotica seedlings.
引文
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[9]Zhu F Y, Chen Y Z, Yan X F. Plant glucosinolate metabolism and sulfur nutrition. Plant Physiology Communications, 2007, 43(6): 1189-1194. 朱凤羽, 陈亚州, 阎秀峰. 植物芥子油苷代谢与硫营养. 植物生理学报, 2007, 43(6): 1189-1194.
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[12]Zheng Q, Liu L, Liu Z P, et al. Comparison of the response of ion distribution in the tissues and cells of the succulent plants Aloe vera and Salicornia europaea to saline stress. Journal of Plant Nutrition and Soil Science, 2009, 172(6): 875.
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[15]Wang Q R, Hock. The research of distribution and redistribution of 35S in Brassica campestris. Chinese Journal of Soil Science, 1998, (1): 29-32. 王庆仁, Hock. 油菜35S分配与再分配的研究. 土壤通报, 1998, (1): 29-32.
[16]Lambers H. SO42- deprivation has an early effect on the content of ribulose-1, 5-bisphosphate carboxylase/oxygenase and photosynthesis in young leaves of wheat. Plant Physiology, 1997, 115(3): 1231-1239.
[17]Massonneau A, Cathala N, Grignon C, et al. Effect of sulphate deficiency on the plasma membrane polypeptide composition of Brassica napus. Journal of Experimental Botany, 1997, 48(306): 93-100.
[18]Zhao F J, Hawkesford M J, Warrilow H G S. Response of two wheat varieties to sulfur addition and diagnosis of sulfur deficiency. Plant Physiology, 1996, 181: 317-327.
[19]Liu M K, Liu G J. Research progress of plant cysteine synthase complex (SAT/OAS-TL). Journal of Anhui Agriculture, 2008, 36(24): 10344-10347. 刘明坤, 刘关君. 植物半胱氨酸合成酶复合体(SAT/OAS-TL)研究进展. 安徽农业科学, 2008, 36(24): 10344-10347.
[20]Kruse J, Kopriva S, Hansch R, et al. Interaction of sulfur and nitrogen nutrition in tobacco (Nicotiana tabacum) plants: Significance of nitrogen source and root nitrate reductase. Plant Biology, 2007, 9: 638-646.
[21]Zhu Y H, Tu N M, Xiao H Q, et al. Effect of sulfur on the growth and physiological and biochemical indices of flue-cures tobacco. Acta Tabacaria Sinica, 2008, 14(4): 28-32. 朱英华, 屠乃美, 肖汉乾, 等. 硫对烤烟生长发育及生理生化指标的影响. 中国烟草学报, 2008, 14(4): 28-32.
[22]Zhu Y J, Li G Q, Guo T C, et al. Effects of sulfur on key enzyme activities involved in nitrogen and sulphur assimilation in flag leaves and grain yield under different nitrogen levels in winter wheat (Triticum aestivum). Acta Agronomica Sinica, 2007, 33(7): 1116-1121. 朱云集, 李国强, 郭天财, 等. 硫对不同氮水平下小麦旗叶氮硫同化关键酶活性及产量的影响. 作物学报, 2007, 33(7): 1116-1121.
[23]Saccomani M, Cacco G, Ferrari G. Effect of nitrogen and/or sulfur deprivation on the uptake and assimilation steps of nitrate and sulfate in maize seedlings. Journal of Plant Nutrition, 1984, 7(7): 1043-1057.
[24]Zhao C, Jiao N Y, Ning T Y, et al. Enzyme activities in nitrogen metabolism of winter wheat and its grain quality under different environmental conditions. Chinese Journal of Applied Ecology, 2006, 17(10): 1866-1870. 赵春, 焦念元, 宁堂原, 等. 不同环境条件下小麦氮代谢关键酶活性及籽粒品质. 应用生态学报, 2006, 17(10): 1866-1870.
[25]Guan Y X, Lin B, Ling B Y. The interactive effects of growth light condition and nitrogen supply on maize (Zea mays) seedling photosynthetic traits and metabolism of carbon and nitrogen. Acta Agronomica Sinica, 2000, 26(6): 806-812. 关义新, 林葆, 凌碧莹. 光、氮及其互作对玉米幼苗叶片光合和碳、氮代谢的影响. 作物学报, 2000, 26(6): 806-812.
[26]Robinson S A, Slade A P, Fox G G, et al. The role of glutamate dehydrogenase in plant nitrogen metabolism. Plant Physiology, 1991, 95: 509-516.
[27]Refouvelet E, Daguin F. Polymorphic glutamate dehydrogenase in lilac vitroplants as revealed by combined preparative IEF and native PAGE: Effect of ammonium deprivation, darkness and atmospheric CO2 enrichment upon isomerization. Physiologia Plantarum, 1999, 105: 199-206.
[28]Lu J W, Chen F, Chen X C, et al. The effect of kalium and nitrogen combined application on the yield and quality of crop. Chinese Journal of Soil Science, 1994, (5): 216-218. 鲁剑巍, 陈防, 陈行春, 等. 钾、硫肥配施对作物产量与品质的影响. 土壤通报, 1994, (5): 216-218.
[29]Zhu F Y. Effect of sulphur deficiency on glucosinolate content in Brassica napus. Harbin: Northeast Forestry University, 2008. 朱凤羽. 硫缺乏对油菜芥子油苷含量的影响. 哈尔滨: 东北林业大学, 2008.
[2]Gao Z C, Zhang X L, Ma X Z. Research advanced in physiological function of element sulphur in plants. Heilongjiang Agriculture Sciences, 2009, (5): 153-155. 高中超, 张喜林, 马星竹. 植物体内硫素的生理功能及作用研究进展. 黑龙江农业科学, 2009, (5): 153-155.
[3]Meng C F, Jiang P K, Cao Z H, et al. Recent progress on transport and assimilation of sulfur in plants. Acta Agriculturae Zhejiangensis, 2011, 23(2): 427-432. 孟赐福, 姜培坤, 曹志洪, 等. 植物体内硫的运输与同化的研究进展. 浙江农业学报, 2011, 23(2): 427-432.
[4]Lewandowska M, Sirko A. Recent advances in understanding plant response to sulfur-deficiency stress. Acta Biochimica Polonica, 2008, 55(3): 457.
[5]Kopriva S, Wiedemann G, Reski R. Sulfate assimilation in basal land plants-what does genomic sequencing tell us? Plant Biology, 2007, 9(5): 556-564.
[6]De Kok L J, Castro A, Durenkamp M, et al. Sulphur in plant physiology. New York: Proceedings No 500, The International Fertiliser Society, 2002: 1-26.
[7]Xie R Z, Dong S T. Progress in sulfur nutrition of plant. Chinese Agricultural Science Bulletin, 2002, 18(2): 65-69. 谢瑞芝, 董树亭. 植物硫素营养研究进展. 中国农学通报, 2002, 18(2): 65-69.
[8]Xu Y, Mou J M, Zhang G Q, et al. Effect of sulphur availability on nitrate accumulation and expression of nitrogen and sulphur assimilation related genes in non-heading Chinese cabbage. Scientia Agricultura Sinica, 2016, 49(11): 2222-2233. 徐瑶, 牟建梅, 张国芹, 等. 施硫对不结球白菜硝酸盐累积及氮硫同化关键基因表达的影响. 中国农业科学, 2016, 49(11): 2222-2233.
[9]Zhu F Y, Chen Y Z, Yan X F. Plant glucosinolate metabolism and sulfur nutrition. Plant Physiology Communications, 2007, 43(6): 1189-1194. 朱凤羽, 陈亚州, 阎秀峰. 植物芥子油苷代谢与硫营养. 植物生理学报, 2007, 43(6): 1189-1194.
[10]Miao Y J, Guan J L, Zeng J L, et al. Effect of nitrogen and sulfur combined application on nutritional components and active components of Isatis indigotica at seedling stage. China Journal of Chinese Materia Medica, 2018, 43(8): 1571-1578. 缪雨静, 关佳莉, 曾佳乐, 等. 氮硫配施对苗期菘蓝中营养物质及活性成分的影响. 中国中药杂志, 2018, 43(8): 1571-1578.
[11]Chinese Pharmacopoeia Commission. Pharmacopoeia of People’s Republic of China. Beijing: China Medical Science Press, 2015: 21, 205. 国家药典委员会. 中华人民共和国药典: 2015版(一部). 北京: 中国医药科技出版社, 2015: 21, 205.
[12]Zheng Q, Liu L, Liu Z P, et al. Comparison of the response of ion distribution in the tissues and cells of the succulent plants Aloe vera and Salicornia europaea to saline stress. Journal of Plant Nutrition and Soil Science, 2009, 172(6): 875.
[13]Wang X K, Zhang W H, Hao Z B, et al. Principles and techniques of plant physiology and biochemistry. Beijing: Higher Education Press, 2006. 王学奎, 章文华, 郝再彬, 等. 植物生理生化实验原理和技术. 北京: 高等教育出版社, 2006.
[14]Friedrich J W, Schrader L E. Sulfur deprivation and nitrogen metabolism in maize seedlings. Plant Physiology, 1978, 61(6): 900-903.
[15]Wang Q R, Hock. The research of distribution and redistribution of 35S in Brassica campestris. Chinese Journal of Soil Science, 1998, (1): 29-32. 王庆仁, Hock. 油菜35S分配与再分配的研究. 土壤通报, 1998, (1): 29-32.
[16]Lambers H. SO42- deprivation has an early effect on the content of ribulose-1, 5-bisphosphate carboxylase/oxygenase and photosynthesis in young leaves of wheat. Plant Physiology, 1997, 115(3): 1231-1239.
[17]Massonneau A, Cathala N, Grignon C, et al. Effect of sulphate deficiency on the plasma membrane polypeptide composition of Brassica napus. Journal of Experimental Botany, 1997, 48(306): 93-100.
[18]Zhao F J, Hawkesford M J, Warrilow H G S. Response of two wheat varieties to sulfur addition and diagnosis of sulfur deficiency. Plant Physiology, 1996, 181: 317-327.
[19]Liu M K, Liu G J. Research progress of plant cysteine synthase complex (SAT/OAS-TL). Journal of Anhui Agriculture, 2008, 36(24): 10344-10347. 刘明坤, 刘关君. 植物半胱氨酸合成酶复合体(SAT/OAS-TL)研究进展. 安徽农业科学, 2008, 36(24): 10344-10347.
[20]Kruse J, Kopriva S, Hansch R, et al. Interaction of sulfur and nitrogen nutrition in tobacco (Nicotiana tabacum) plants: Significance of nitrogen source and root nitrate reductase. Plant Biology, 2007, 9: 638-646.
[21]Zhu Y H, Tu N M, Xiao H Q, et al. Effect of sulfur on the growth and physiological and biochemical indices of flue-cures tobacco. Acta Tabacaria Sinica, 2008, 14(4): 28-32. 朱英华, 屠乃美, 肖汉乾, 等. 硫对烤烟生长发育及生理生化指标的影响. 中国烟草学报, 2008, 14(4): 28-32.
[22]Zhu Y J, Li G Q, Guo T C, et al. Effects of sulfur on key enzyme activities involved in nitrogen and sulphur assimilation in flag leaves and grain yield under different nitrogen levels in winter wheat (Triticum aestivum). Acta Agronomica Sinica, 2007, 33(7): 1116-1121. 朱云集, 李国强, 郭天财, 等. 硫对不同氮水平下小麦旗叶氮硫同化关键酶活性及产量的影响. 作物学报, 2007, 33(7): 1116-1121.
[23]Saccomani M, Cacco G, Ferrari G. Effect of nitrogen and/or sulfur deprivation on the uptake and assimilation steps of nitrate and sulfate in maize seedlings. Journal of Plant Nutrition, 1984, 7(7): 1043-1057.
[24]Zhao C, Jiao N Y, Ning T Y, et al. Enzyme activities in nitrogen metabolism of winter wheat and its grain quality under different environmental conditions. Chinese Journal of Applied Ecology, 2006, 17(10): 1866-1870. 赵春, 焦念元, 宁堂原, 等. 不同环境条件下小麦氮代谢关键酶活性及籽粒品质. 应用生态学报, 2006, 17(10): 1866-1870.
[25]Guan Y X, Lin B, Ling B Y. The interactive effects of growth light condition and nitrogen supply on maize (Zea mays) seedling photosynthetic traits and metabolism of carbon and nitrogen. Acta Agronomica Sinica, 2000, 26(6): 806-812. 关义新, 林葆, 凌碧莹. 光、氮及其互作对玉米幼苗叶片光合和碳、氮代谢的影响. 作物学报, 2000, 26(6): 806-812.
[26]Robinson S A, Slade A P, Fox G G, et al. The role of glutamate dehydrogenase in plant nitrogen metabolism. Plant Physiology, 1991, 95: 509-516.
[27]Refouvelet E, Daguin F. Polymorphic glutamate dehydrogenase in lilac vitroplants as revealed by combined preparative IEF and native PAGE: Effect of ammonium deprivation, darkness and atmospheric CO2 enrichment upon isomerization. Physiologia Plantarum, 1999, 105: 199-206.
[28]Lu J W, Chen F, Chen X C, et al. The effect of kalium and nitrogen combined application on the yield and quality of crop. Chinese Journal of Soil Science, 1994, (5): 216-218. 鲁剑巍, 陈防, 陈行春, 等. 钾、硫肥配施对作物产量与品质的影响. 土壤通报, 1994, (5): 216-218.
[29]Zhu F Y. Effect of sulphur deficiency on glucosinolate content in Brassica napus. Harbin: Northeast Forestry University, 2008. 朱凤羽. 硫缺乏对油菜芥子油苷含量的影响. 哈尔滨: 东北林业大学, 2008.