种植密度对烤烟光合特性日变化及其主要化学成分的影响
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摘要
[目的]探讨种植密度对烤烟光合特性日变化及其主要化学成分的影响,为协调烤烟的碳、氮代谢,优化烤后烟叶化学成分的协调性提供理论依据。[方法]以烤烟(Nicotiana tabacum L.)品种‘K326’为试验材料,设置5个种植密度处理,即最小密度处理(T1,14 200株·hm~(-2))、较小密度处理(T2,15 400株·hm~(-2))、中等密度处理(T3,16 800株·hm~(-2))、较大密度处理(T4,18 500株·hm~(-2))、最大密度处理(T5,20 500株·hm~(-2)),研究种植密度对烤烟光合特性和常规化学成分的影响。[结果]除最大密度处理外,其他处理烤烟净光合速率均呈双峰曲线,峰值出现在11:00和15:00;与最小密度处理相比,较小密度处理烤烟中部叶净光合速率、气孔导度、蒸腾速率分别提高14.59%、13.11%、3.25%,13:00之后胞间CO_2浓度回升加快,气孔限制值逐渐下降,说明气孔限制得到有效缓解,有利于延长烟叶光合功能时间,烤后烟叶两糖比升高了4.40%,成熟度提高;最大密度处理的烤烟中部叶13:00之后净光合速率下降92.30%,胞间CO_2浓度升高18.52%,气孔限制值下降17.50%,说明随着光合作用的进行,烟叶非气孔限制加剧,光合效率低,有效光合时间短,因此不利于干物质积累;净光合速率、气孔导度、蒸腾速率均与氮碱比呈极显著正相关,蒸腾速率与总糖呈显著正相关。[结论]与最大密度处理相比,较大和中等密度处理有利于优化烟株形态,协调烟叶光合速率,提高烤烟成熟度及碳、氮代谢协调性,实现优质烟叶生产。
[Objectives]The objectives of this study were to investigate the effects of plant density on diurnal variation of photosynthetic characteristics and main chemical components of flue-cured tobacco,and to provide a theoretical basis for coordinating the carbon and nitrogen metabolism of flue-cured tobacco and optimizing the coordination of chemical components in tobacco leaves. [Methods]The Nicotiana tabacum L. variety'K326'was planted at minimum density treatment(T1,14 200 plants·hm~(-2)),small density treatment(T2,15 400 plants·hm~(-2)),medium density treatment(T3,16 800 plants·hm~(-2)),larger density treatment(T4,18 500 plants·hm~(-2)),and maximum density treatment(T5,20 500 plants·hm~(-2)). The effects of densification on photosynthetic characteristics and chemical composition of flue-cured tobacco were studied. [Results]Compared with the minimum density treatment,the net photosynthetic rate,stomatal conductance and transpiration rate of the middle leaves with the small density treatment increased by 14.59%,13.11% and 3.25%,respectively.Intercellular CO_2concentration of the middle leaves with the small density treatment rose faster and stomatal limit value decreased gradually after 13:00,indicating that stomatal limit was effectively alleviated,which was conducive to prolonging the photosynthetic function time of tobacco leaves. The ratio of two sugars in tobacco leaves treated with the small density treatment increased by 4.40% compared with the minimum density treatment,and this means that the maturity was improved. After 13:00,net photosynthetic rate decreased by 92.30%,intercellular CO_2 concentration increased by 18.52%,and stomatal limit value decreased by 17.50%. The results showed that the non-stomatal restriction increased,the photosynthetic efficiency was low and the effective photosynthetic time was short,and it was not conducive to dry matter accumulation. Net photosynthetic rate,stomatal conductance and transpiration rate were significantly positively correlated with the ratio of nitrogen to alkali,and transpiration rate was significantly positively correlated with total sugar. [Conclusions]Compared with the maximum density,the treatment with larger and medium density was beneficial to optimizing the tobacco plant morphology,coordinated the photosynthetic rate of tobacco leaves,improved the maturity of flue-cured tobacco and the coordination of carbon and nitrogen metabolism,and achieved the production of high-quality tobacco leaves.
[Objectives]The objectives of this study were to investigate the effects of plant density on diurnal variation of photosynthetic characteristics and main chemical components of flue-cured tobacco,and to provide a theoretical basis for coordinating the carbon and nitrogen metabolism of flue-cured tobacco and optimizing the coordination of chemical components in tobacco leaves. [Methods]The Nicotiana tabacum L. variety'K326'was planted at minimum density treatment(T1,14 200 plants·hm~(-2)),small density treatment(T2,15 400 plants·hm~(-2)),medium density treatment(T3,16 800 plants·hm~(-2)),larger density treatment(T4,18 500 plants·hm~(-2)),and maximum density treatment(T5,20 500 plants·hm~(-2)). The effects of densification on photosynthetic characteristics and chemical composition of flue-cured tobacco were studied. [Results]Compared with the minimum density treatment,the net photosynthetic rate,stomatal conductance and transpiration rate of the middle leaves with the small density treatment increased by 14.59%,13.11% and 3.25%,respectively.Intercellular CO_2concentration of the middle leaves with the small density treatment rose faster and stomatal limit value decreased gradually after 13:00,indicating that stomatal limit was effectively alleviated,which was conducive to prolonging the photosynthetic function time of tobacco leaves. The ratio of two sugars in tobacco leaves treated with the small density treatment increased by 4.40% compared with the minimum density treatment,and this means that the maturity was improved. After 13:00,net photosynthetic rate decreased by 92.30%,intercellular CO_2 concentration increased by 18.52%,and stomatal limit value decreased by 17.50%. The results showed that the non-stomatal restriction increased,the photosynthetic efficiency was low and the effective photosynthetic time was short,and it was not conducive to dry matter accumulation. Net photosynthetic rate,stomatal conductance and transpiration rate were significantly positively correlated with the ratio of nitrogen to alkali,and transpiration rate was significantly positively correlated with total sugar. [Conclusions]Compared with the maximum density,the treatment with larger and medium density was beneficial to optimizing the tobacco plant morphology,coordinated the photosynthetic rate of tobacco leaves,improved the maturity of flue-cured tobacco and the coordination of carbon and nitrogen metabolism,and achieved the production of high-quality tobacco leaves.
引文
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[8] 常爱霞,张建平,杜咏梅,等.烤烟香型相关化学成分主导的不同产区烟叶聚类分析[J].中国烟草学报,2010,16(2):14-19.Chang A X,Zhang J P,Du Y M,et al.Cluster analysis of flue-cured tobacco leaves from different growing areas according to the chemical components correlating with aroma types[J].Acta Tabacaria Sinica,2010,16(2):14-19(in Chinese with English abstract).
[9] 李旭华,何传国,陈建军,等.广东浓香型特色烟叶关键生产技术理论与应用[M].广州:华南理工大学出版社,2011:12.Li X H,He C G,Chen J J,et al.Theory and Application of Key Production Technology of Guangdong Luzhou-flavored Characteristic Tobacco Leaves[M].Guangzhou:South China University of Technology Press,2011:12(in Chinese).
[10] 许海良,赵会杰,蒲文宣,等.种植密度对烟草冠层辐射和群体光合作用的影响[J].福建农业学报,2017,32(3):253-257.Xu H L,Zhao H J,Pu W X,et al.Effect of planting density on light transmittance and canopy photosynthesis of flue-cured tobacco plants[J].Fujian Journal of Agricultural Sciences,2017,32(3):253-257(in Chinese with English abstract).
[11] 王瑞,刘国顺,倪国仕,等.种植密度对烤烟不同部位叶片光合特性及其同化物积累的影响[J].作物学报,2009,35(12):2288-2295.Wang R,Liu G S,Ni G S,et al.Effects of planting density on photosynthetic characteristics and assimilate accumulation of leaves in different positions in flue-cured tobacco[J].Acta Agronomica Sinica,2009,35(12):2288-2295(in Chinese with English abstract).
[12] 叶卫国,雷佳,李淮源,等.减氮增密对烤烟光合特性及烟叶结构的影响[J].南京农业大学学报,2018,41(5):817-824.DOI:10.7685/jnau.201802007.Ye W G,Lei J,Li H Y,et al.Effects of nitrogen-saving and density-increasing on photosynthetic characteristics and leaf structure of flue-cured tobacco[J].Journal of Nanjing Agricultural University,2018,41(5):817-824(in Chinese with English abstract).
[13] 贾方方.不同种植密度烟草叶面积指数的高光谱估测模型[J].中国烟草科学,2017,38(4):37-43.Jia F F.Estimating model for leaf area index of tobacco via hyperspectral reflectance at different planting densities[J].Chinese Tobacco Science,2017,38(4):37-43(in Chinese with English abstract).
[14] 肖金香,刘正和,王燕,等.气候生态因素对烤烟产量与品质的影响及植烟措施研究[J].中国生态农业学报,2003,11(4):158-160.Xiao J X,Liu Z H,Wang Y,et al.The influence of climatic ecology factors on the yield and quality of flue-cured tobacco and the countermeasures of planting tobacco in Jiangxi[J].Chinese Journal of Eco-Agriculture,2003,11(4):158-160(in Chinese with English abstract).
[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,2006:186-199.Li H S.Principles and Techniques of Plant Physiological and Biochemical Experiments[M].Beijing:Higher Education Press,2006:186-199(in Chinese).
[16] 王瑞新.烟草化学[M].北京:中国农业出版社,2003:250-286.Wang R X.Tobacco Chemistry[M].Beijing:China Agriculture Press,2003:250-286(in Chinese).
[17] 张振文,林立铭,余厚美,等.温度对木薯叶片叶绿素荧光及 Rubisco 酶的影响[J].江西农业学报,2017,29(1):1-5.Zhang Z W,Lin L M,Yu H M,et al.Effects of temperature on chlorophyll fluorescence parameters and rubisco activity in cassava leaves[J].Acta Agriculturae Jiangxi,2017,29(1):1-5(in Chinese with English abstract).
[18] 邓世媛,谢晋,周立非,等.精量施氮对烤烟成熟期叶片碳氮代谢的影响[J].烟草科技,2013,46(9):76-81.Deng S Y,Xie J,Zhou L F,et al.Effects of precise nitrogen application on carbon and nitrogen metabolism of flue-cured tobacco leaves at maturity stage[J].Tobacco Science & Technology,2013,46(9):76-81(in Chinese with English abstract).
[19] 魏彬,陈建军,吕永华,等.不同调控措施下烤烟开花时间与碳氮代谢关系的研究[J].中国烟草科学,2007,28(1):14-17.Wei B,Chen J J,Lü Y H,et al.Effects of various treatments on flowering time and C/N metabolism of flue-cured tobacco[J].Chinese Tobacco Science,2007,28(1):14-17(in Chinese with English abstract).
[20] 许自成,黄平俊,卢秀萍,等.不同覆盖措施对大田烤烟叶片淀粉积累的影响[J].西北农林科技大学学报(自然科学版),2006,34(5):77-82.Xu Z C,Huang P J,Lu X P,et al.Effects of different mulching measures on starch accumulation in flue-cured tobacco leaves during field growth[J].Journal of Northwest A&F University(Natural Science Edition),2006,34(5):77-82(in Chinese with English abstract).
[21] 宫长荣.烟草调制学[M].北京:中国农业出版社,2003:156-157.Gong C R.Tobacco Curing[M].Beijing:China Agriculture Press,2003:156-157(in Chinese).
[22] 曹阳,钟俊周,文国宇,等.种植密度对烤烟生长发育及烘烤特性的影响[J].安徽农业科学,2018,46(28):35-37.Cao Y,Zhong J Z,Wen G Y,et al.Effects of planting density on the tobacco growth development and curing characteristics[J].Journal of Anhui Agricultural Sciences,2018,46(28):35-37(in Chinese with English abstract).
[2] 胡近近,钟俊周,陈君豪,等.不同供氮形态对烤烟烘烤特性的影响[J].华北农学报,2017,32(3):174-181.Hu J J,Zhong J Z,Chen J H,et al.Effects of different nitrogen forms on curing characteristics in flue-cured tobacco[J].Acta Agriculturae Boreali-Sinica,2017,32(3):174-181(in Chinese with English abstract).
[3] Tang H,Hu Y Y,Yu W W,et al.Growth,photosynthetic and physiological responses of Torreya grandis seedlings to varied light environments[J].Trees,2015,29(4):1011-1022.
[4] 高升.施氮量与种植密度对烤烟品种K326烟株生长和烟叶品质的影响[D].重庆:西南大学,2016.Gao S.Effects of nitrogen application rate and planting density on growth and quality of flue-cured tobacco K326[D].Chongqing:Southwest University,2016(in Chinese with English abstract).
[5] 王传义.不同烤烟品种烘烤特性研究[D].北京:中国农业科学院,2008.Wang C Y.Study on the leaf flue curing characteristics of different varieties of flue-cured tobacco[D].Beijing:Chinese Academy of Agricultural Sciences,2008(in Chinese with English abstract).
[6] Karaivazoglou N A,Tsotsolis N C,Tsadilas C D.Influence of liming and form of nitrogen fertilizer on nutrient uptake,growth,yield,and quality of Virginia(flue-cured)tobacco[J].Field Crops Research,2007,100(1):52-60.
[7] 秦松,王正银,石俊雄,等.贵州省不同香气类型烟叶质量特征研究[J].中国农业科学,2006,39(11):2319-2326.Qin S,Wang Z Y,Shi J X,et al.Study on quality character of leaf for different types of aroma in Guizhou Province[J].Scientia Agricultura Sinica,2006,39(11):2319-2326(in Chinese with English abstract).
[8] 常爱霞,张建平,杜咏梅,等.烤烟香型相关化学成分主导的不同产区烟叶聚类分析[J].中国烟草学报,2010,16(2):14-19.Chang A X,Zhang J P,Du Y M,et al.Cluster analysis of flue-cured tobacco leaves from different growing areas according to the chemical components correlating with aroma types[J].Acta Tabacaria Sinica,2010,16(2):14-19(in Chinese with English abstract).
[9] 李旭华,何传国,陈建军,等.广东浓香型特色烟叶关键生产技术理论与应用[M].广州:华南理工大学出版社,2011:12.Li X H,He C G,Chen J J,et al.Theory and Application of Key Production Technology of Guangdong Luzhou-flavored Characteristic Tobacco Leaves[M].Guangzhou:South China University of Technology Press,2011:12(in Chinese).
[10] 许海良,赵会杰,蒲文宣,等.种植密度对烟草冠层辐射和群体光合作用的影响[J].福建农业学报,2017,32(3):253-257.Xu H L,Zhao H J,Pu W X,et al.Effect of planting density on light transmittance and canopy photosynthesis of flue-cured tobacco plants[J].Fujian Journal of Agricultural Sciences,2017,32(3):253-257(in Chinese with English abstract).
[11] 王瑞,刘国顺,倪国仕,等.种植密度对烤烟不同部位叶片光合特性及其同化物积累的影响[J].作物学报,2009,35(12):2288-2295.Wang R,Liu G S,Ni G S,et al.Effects of planting density on photosynthetic characteristics and assimilate accumulation of leaves in different positions in flue-cured tobacco[J].Acta Agronomica Sinica,2009,35(12):2288-2295(in Chinese with English abstract).
[12] 叶卫国,雷佳,李淮源,等.减氮增密对烤烟光合特性及烟叶结构的影响[J].南京农业大学学报,2018,41(5):817-824.DOI:10.7685/jnau.201802007.Ye W G,Lei J,Li H Y,et al.Effects of nitrogen-saving and density-increasing on photosynthetic characteristics and leaf structure of flue-cured tobacco[J].Journal of Nanjing Agricultural University,2018,41(5):817-824(in Chinese with English abstract).
[13] 贾方方.不同种植密度烟草叶面积指数的高光谱估测模型[J].中国烟草科学,2017,38(4):37-43.Jia F F.Estimating model for leaf area index of tobacco via hyperspectral reflectance at different planting densities[J].Chinese Tobacco Science,2017,38(4):37-43(in Chinese with English abstract).
[14] 肖金香,刘正和,王燕,等.气候生态因素对烤烟产量与品质的影响及植烟措施研究[J].中国生态农业学报,2003,11(4):158-160.Xiao J X,Liu Z H,Wang Y,et al.The influence of climatic ecology factors on the yield and quality of flue-cured tobacco and the countermeasures of planting tobacco in Jiangxi[J].Chinese Journal of Eco-Agriculture,2003,11(4):158-160(in Chinese with English abstract).
[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,2006:186-199.Li H S.Principles and Techniques of Plant Physiological and Biochemical Experiments[M].Beijing:Higher Education Press,2006:186-199(in Chinese).
[16] 王瑞新.烟草化学[M].北京:中国农业出版社,2003:250-286.Wang R X.Tobacco Chemistry[M].Beijing:China Agriculture Press,2003:250-286(in Chinese).
[17] 张振文,林立铭,余厚美,等.温度对木薯叶片叶绿素荧光及 Rubisco 酶的影响[J].江西农业学报,2017,29(1):1-5.Zhang Z W,Lin L M,Yu H M,et al.Effects of temperature on chlorophyll fluorescence parameters and rubisco activity in cassava leaves[J].Acta Agriculturae Jiangxi,2017,29(1):1-5(in Chinese with English abstract).
[18] 邓世媛,谢晋,周立非,等.精量施氮对烤烟成熟期叶片碳氮代谢的影响[J].烟草科技,2013,46(9):76-81.Deng S Y,Xie J,Zhou L F,et al.Effects of precise nitrogen application on carbon and nitrogen metabolism of flue-cured tobacco leaves at maturity stage[J].Tobacco Science & Technology,2013,46(9):76-81(in Chinese with English abstract).
[19] 魏彬,陈建军,吕永华,等.不同调控措施下烤烟开花时间与碳氮代谢关系的研究[J].中国烟草科学,2007,28(1):14-17.Wei B,Chen J J,Lü Y H,et al.Effects of various treatments on flowering time and C/N metabolism of flue-cured tobacco[J].Chinese Tobacco Science,2007,28(1):14-17(in Chinese with English abstract).
[20] 许自成,黄平俊,卢秀萍,等.不同覆盖措施对大田烤烟叶片淀粉积累的影响[J].西北农林科技大学学报(自然科学版),2006,34(5):77-82.Xu Z C,Huang P J,Lu X P,et al.Effects of different mulching measures on starch accumulation in flue-cured tobacco leaves during field growth[J].Journal of Northwest A&F University(Natural Science Edition),2006,34(5):77-82(in Chinese with English abstract).
[21] 宫长荣.烟草调制学[M].北京:中国农业出版社,2003:156-157.Gong C R.Tobacco Curing[M].Beijing:China Agriculture Press,2003:156-157(in Chinese).
[22] 曹阳,钟俊周,文国宇,等.种植密度对烤烟生长发育及烘烤特性的影响[J].安徽农业科学,2018,46(28):35-37.Cao Y,Zhong J Z,Wen G Y,et al.Effects of planting density on the tobacco growth development and curing characteristics[J].Journal of Anhui Agricultural Sciences,2018,46(28):35-37(in Chinese with English abstract).