种植密度对烤烟叶片碳同化能力及同化产物分配的影响
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摘要
以烤烟品种K326为材料,设置D1(16 529株/hm2)、D2(18 182株/hm2)、D3(20 202株/hm2)3个种植密度处理,于圆顶期对各处理烟株上、中、下3个部位叶片进行13C同位素标记,研究了烤烟不同部位叶片的碳同化能力差异和同化产物的分配特点,并探讨了种植密度对烤烟叶片碳同化能力差异和同化产物分配的调节效应。结果表明,烤烟不同部位叶片的13C同化能力表现为中部叶>上部叶>下部叶;同一叶片不同区段13C同化能力大小依次为叶中部>叶基部>叶尖部。上部叶的13C同化产物平均有19.71%运往根系,42.22%运往茎秆,38.07%留在叶片且大部分留在被标记的上部叶;中部叶的13C同化产物有33.42%运往根系,26.00%运往茎秆,40.58%留在叶片且大部分留在被标记的中部叶;下部叶的13C同化产物有33.67%运往根系,23.72%运往茎秆,42.61%留在叶片且大部分留在被标记的下部叶。上部叶的13C同化产物向茎秆的中部供应较多,中部叶和下部叶的13C同化产物向茎秆基部分配较多。随着种植密度增大,叶片的碳同化能力降低,向根、茎分配的同化产物量减少,尤其是在较高的种植密度(D3)下,各部位叶片的碳同化能力显著降低。综上,烟叶产量和品质形成的物质基础主要来源于烟叶生长期间自身的同化作用,种植密度对烤烟叶片的13C同化能力具有显著影响,在设置试验条件下,18 182株/hm2左右的种植密度有利于提高烤烟叶片碳同化能力和协调同化产物的运输分配。
In present study,13C labelling method was used to evaluate the differences in carbon assimilation ability of leaves at different positions of flue-cured tobacco plants,the distribution characteristics of photosynthate,and the regulating effect of planting density. Three different treatments of planting density were designed and denoted as D1( 16 529 plants/ha),D2( 18 182 plants/ha) and D3( 20 202 plants/ha). The leaves at upper,middle and lower positions of tobacco plants were labeled with13 CO2 respectively at round top stage. As shown in the results,at the same flue-cured tobacco plant,the highest13 C assimilation capability was observed on middle leaves,followed by the upper and the lower leaves. In comparison with different areas of the same leaf,the highest capability of carbon assimilationwas in the middle section,followed by the base and the top section. Averagely,19. 71% of13 C assimilation products of upper leaves were transported to root system,42. 22% to stem,and 38. 07% were left in leaves and most of them were left in upper labelled leaves. 33. 42% of13 C assimilation products of middle leaves were transported to root system,26. 00% to stem,and 40. 58% were left in leaves and most of them were left in middle labelled leaves. 33. 67% of13 C assimilation products made by lower leaves were transported to root system,23. 72% to stem,and 42. 61% were left in leaves and most of them were left in lower labelled leaves. In terms of assimilation product allocated to the stem,photosynthates of upper leaves were mainly distributed to middle part of stem,and photosynthates of middle and lower leaves were mainly distributed to base part of stem. The carbon assimilation capacity of leaves and the amount of assimilation product transported to root and stem both decreased along with the increase of planting density. Under the high planting density of D3,carbon assimilation capacity of leaves at all positions declined significantly. In conclusion,substance basis for the tobacco yield and quality mainly originates from the carbon assimilation by themselves during the growth of leaves. The planting density has a significant impact on the13 C assimilative capacity of flue-cured tobacco leaves. Under this experimental conditions,the planting density around 18 182 plants/ha is conducive to improving the carbon assimilation capacity and coordinating the assimilation product distribution in flue-cured tobacco.
In present study,13C labelling method was used to evaluate the differences in carbon assimilation ability of leaves at different positions of flue-cured tobacco plants,the distribution characteristics of photosynthate,and the regulating effect of planting density. Three different treatments of planting density were designed and denoted as D1( 16 529 plants/ha),D2( 18 182 plants/ha) and D3( 20 202 plants/ha). The leaves at upper,middle and lower positions of tobacco plants were labeled with13 CO2 respectively at round top stage. As shown in the results,at the same flue-cured tobacco plant,the highest13 C assimilation capability was observed on middle leaves,followed by the upper and the lower leaves. In comparison with different areas of the same leaf,the highest capability of carbon assimilationwas in the middle section,followed by the base and the top section. Averagely,19. 71% of13 C assimilation products of upper leaves were transported to root system,42. 22% to stem,and 38. 07% were left in leaves and most of them were left in upper labelled leaves. 33. 42% of13 C assimilation products of middle leaves were transported to root system,26. 00% to stem,and 40. 58% were left in leaves and most of them were left in middle labelled leaves. 33. 67% of13 C assimilation products made by lower leaves were transported to root system,23. 72% to stem,and 42. 61% were left in leaves and most of them were left in lower labelled leaves. In terms of assimilation product allocated to the stem,photosynthates of upper leaves were mainly distributed to middle part of stem,and photosynthates of middle and lower leaves were mainly distributed to base part of stem. The carbon assimilation capacity of leaves and the amount of assimilation product transported to root and stem both decreased along with the increase of planting density. Under the high planting density of D3,carbon assimilation capacity of leaves at all positions declined significantly. In conclusion,substance basis for the tobacco yield and quality mainly originates from the carbon assimilation by themselves during the growth of leaves. The planting density has a significant impact on the13 C assimilative capacity of flue-cured tobacco leaves. Under this experimental conditions,the planting density around 18 182 plants/ha is conducive to improving the carbon assimilation capacity and coordinating the assimilation product distribution in flue-cured tobacco.
引文
[1]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.
[2]时向东,朱命阳,赵会纳,等.种植密度对烤烟叶片生育期光合特性的影响[J].中国烟草学报,2012,18(6):38-42.
[3]汪耀富,孙德梅,韩富根,等.密度和地膜覆盖对烟田冠层生理特性和土壤水分利用效率的影响[J].烟草科技,2003(12):27-30.
[4]李海平,朱列书,黄魏魏,等.种植密度对烟田环境、烤烟农艺性状及产量质量的影响研究进展[J].作物研究,2008,22(5):489-490.
[5]毛家伟,张翔,王宏,等.种植密度和氮用量对烟叶光合特性和产量质量的影响[J].干旱地区农业研究,2012,35(5):66-70.
[6]沈其荣,殷士学,杨超光,等.13C标记技术在土壤和植物营养研究中的应用[J].植物营养与肥料学报,2000,6(1):98-105.
[7]马晔,刘锦春.δ13C在植物生态学研究中的应用[J].西北植物学报,2013,33(7):1492-1500.
[8]Liu R,Clapp C E,Cheng H H.Usefulness of the carbon-13 tracer technique for characerizing terrestrial carbon pools[J].Nutrient Cycling in Agroecosystems,1997,49(1/2/3):261-266.
[9]Balesdent J,Mariotti A,Boisgontier D.Effect of tillage on soil organic matter carbon mineralization estimated from13C abundance in maize fields[J].Journal of Soil Science,1990,41:587-596.
[10]Bernoux M,Cerri C C,Neill C,et al.The use of stable carbon isotopes for estimating soil organic matter turnover rates[J].Geoderma,1998,82(1/3):43-58.
[11]Gebrekirstos A,Van Noordwijk M,Neufeldt H,et al.Relationships of stable carbon isotopes,plant water potential and growth:An approach to asses water use efficiency and growth strategies of dry land agroforestry species[J].Trees,2011,25(1):95-102.
[12]Miller J M,Williams R J,Farquhar G D.Carbon isotope discrimination by a sequence of Eucalyptus species along a subcontinental rainfall gradient in Australia[J].Functional Ecology,2001,15:222-232.
[13]何春霞,李吉跃,孟平,等.树木叶片稳定碳同位素分馏对环境梯度的响应[J].生态学报,2010,30(14):3828-3838.
[14]齐鑫,王敬国.应用13C脉冲标记方法研究不同施氮量对冬小麦净光合碳分配及其向地下输入的影响[J].农业环境科学学报,2008,27(6):2524-2530.
[15]赵隽,董树亭,刘鹏,等.有机无机肥长期定位配施对冬小麦13C同化物分配及产量的影响[J].核农学报,2016,30(9):1805-1814.
[16]何敏毅,孟凡乔,史雅娟,等.用13C脉冲标记法研究玉米光合碳分配及其向地下的输入[J].环境科学,2008,29(2):446-453.
[17]安婷婷,汪景宽,李双异,等.用13C脉冲标记方法研究施肥与地膜覆盖对玉米光合碳分配的影响[J].土壤学报,2013,50(5):948-955.
[18]Lu Y H,Watanabe A,Kimura M.Input and distribution of photosynthesized carbon in a flooded rice soil[J].Global Biogeochemical Cycles,2002,16(4):1085-1093.
[19]Lu Y H,Watanabe A,Kimura M.Carbon dynamics of rhizodeposits,root-and shoot-residues in a rice soil[J].Soil Biology&Biochemistry,2003,35(9):1223-1230.
[20]李晶,姜远茂,魏绍冲,等.富士苹果秋梢连续摘心对13C和15N利用、分配的影响[J].园艺学报,2012,39(11):2238-2244.
[21]门永阁,任饴华,许海港,等.苹果树不同部位新梢叶片13C同化物的去向[J].园艺学报,2014,41(6):1063-1068.
[22]付莹,姜远茂,张世忠,等.短截修剪程度对‘红灯’甜樱桃13C和15N分配利用的影响[J].园艺学报,2015,42(1):104-110.
[23]朱列书,赵松义,李伟.烟草不同基因型的光合特性研究[J].中国烟草科学,2006,27(1):5-7.
[24]王艳丽,刘国顺,李海江,等.烤烟不同部位叶片的光合特性研究[J].中国农学通报,2014,30(10):92-98.
[25]闫川,洪晓富,阮关海,等.大穗型水稻13C光合产物的积累与分配[J].核农学报,2014,28(7):1282-1287.
[26]马冬云,郭天财,王晨阳,等.施氮量对冬小麦灌浆期光合产物积累、转运及分配的影响[J].作物学报,2008,34(6):1027-1033.
[27]姜东,于振文,李永庚,等.高产小麦营养器官临时贮存物质积运及其对粒重的贡献[J].作物学报,2003,29(1):31-36.
[28]杨俊峰,龚月桦,王俊儒,等.旱地覆膜对冬小麦花后14C-同化物转运分配的影响[J].核农学报,2007,21(1):70-74.
[29]于凤义,张萍,周洪杰.玉米功能叶片碳同化物在果穗不同部位籽粒中的分配[J].核农学报,1994,10(1):35-38.
[30]张立军,梁宗锁.植物生理学[M].北京:科学出版社,2012:184-186.
[31]李合生.现代植物生理学[M].3版.北京:高等教育出版社,2012:186-190.
[2]时向东,朱命阳,赵会纳,等.种植密度对烤烟叶片生育期光合特性的影响[J].中国烟草学报,2012,18(6):38-42.
[3]汪耀富,孙德梅,韩富根,等.密度和地膜覆盖对烟田冠层生理特性和土壤水分利用效率的影响[J].烟草科技,2003(12):27-30.
[4]李海平,朱列书,黄魏魏,等.种植密度对烟田环境、烤烟农艺性状及产量质量的影响研究进展[J].作物研究,2008,22(5):489-490.
[5]毛家伟,张翔,王宏,等.种植密度和氮用量对烟叶光合特性和产量质量的影响[J].干旱地区农业研究,2012,35(5):66-70.
[6]沈其荣,殷士学,杨超光,等.13C标记技术在土壤和植物营养研究中的应用[J].植物营养与肥料学报,2000,6(1):98-105.
[7]马晔,刘锦春.δ13C在植物生态学研究中的应用[J].西北植物学报,2013,33(7):1492-1500.
[8]Liu R,Clapp C E,Cheng H H.Usefulness of the carbon-13 tracer technique for characerizing terrestrial carbon pools[J].Nutrient Cycling in Agroecosystems,1997,49(1/2/3):261-266.
[9]Balesdent J,Mariotti A,Boisgontier D.Effect of tillage on soil organic matter carbon mineralization estimated from13C abundance in maize fields[J].Journal of Soil Science,1990,41:587-596.
[10]Bernoux M,Cerri C C,Neill C,et al.The use of stable carbon isotopes for estimating soil organic matter turnover rates[J].Geoderma,1998,82(1/3):43-58.
[11]Gebrekirstos A,Van Noordwijk M,Neufeldt H,et al.Relationships of stable carbon isotopes,plant water potential and growth:An approach to asses water use efficiency and growth strategies of dry land agroforestry species[J].Trees,2011,25(1):95-102.
[12]Miller J M,Williams R J,Farquhar G D.Carbon isotope discrimination by a sequence of Eucalyptus species along a subcontinental rainfall gradient in Australia[J].Functional Ecology,2001,15:222-232.
[13]何春霞,李吉跃,孟平,等.树木叶片稳定碳同位素分馏对环境梯度的响应[J].生态学报,2010,30(14):3828-3838.
[14]齐鑫,王敬国.应用13C脉冲标记方法研究不同施氮量对冬小麦净光合碳分配及其向地下输入的影响[J].农业环境科学学报,2008,27(6):2524-2530.
[15]赵隽,董树亭,刘鹏,等.有机无机肥长期定位配施对冬小麦13C同化物分配及产量的影响[J].核农学报,2016,30(9):1805-1814.
[16]何敏毅,孟凡乔,史雅娟,等.用13C脉冲标记法研究玉米光合碳分配及其向地下的输入[J].环境科学,2008,29(2):446-453.
[17]安婷婷,汪景宽,李双异,等.用13C脉冲标记方法研究施肥与地膜覆盖对玉米光合碳分配的影响[J].土壤学报,2013,50(5):948-955.
[18]Lu Y H,Watanabe A,Kimura M.Input and distribution of photosynthesized carbon in a flooded rice soil[J].Global Biogeochemical Cycles,2002,16(4):1085-1093.
[19]Lu Y H,Watanabe A,Kimura M.Carbon dynamics of rhizodeposits,root-and shoot-residues in a rice soil[J].Soil Biology&Biochemistry,2003,35(9):1223-1230.
[20]李晶,姜远茂,魏绍冲,等.富士苹果秋梢连续摘心对13C和15N利用、分配的影响[J].园艺学报,2012,39(11):2238-2244.
[21]门永阁,任饴华,许海港,等.苹果树不同部位新梢叶片13C同化物的去向[J].园艺学报,2014,41(6):1063-1068.
[22]付莹,姜远茂,张世忠,等.短截修剪程度对‘红灯’甜樱桃13C和15N分配利用的影响[J].园艺学报,2015,42(1):104-110.
[23]朱列书,赵松义,李伟.烟草不同基因型的光合特性研究[J].中国烟草科学,2006,27(1):5-7.
[24]王艳丽,刘国顺,李海江,等.烤烟不同部位叶片的光合特性研究[J].中国农学通报,2014,30(10):92-98.
[25]闫川,洪晓富,阮关海,等.大穗型水稻13C光合产物的积累与分配[J].核农学报,2014,28(7):1282-1287.
[26]马冬云,郭天财,王晨阳,等.施氮量对冬小麦灌浆期光合产物积累、转运及分配的影响[J].作物学报,2008,34(6):1027-1033.
[27]姜东,于振文,李永庚,等.高产小麦营养器官临时贮存物质积运及其对粒重的贡献[J].作物学报,2003,29(1):31-36.
[28]杨俊峰,龚月桦,王俊儒,等.旱地覆膜对冬小麦花后14C-同化物转运分配的影响[J].核农学报,2007,21(1):70-74.
[29]于凤义,张萍,周洪杰.玉米功能叶片碳同化物在果穗不同部位籽粒中的分配[J].核农学报,1994,10(1):35-38.
[30]张立军,梁宗锁.植物生理学[M].北京:科学出版社,2012:184-186.
[31]李合生.现代植物生理学[M].3版.北京:高等教育出版社,2012:186-190.