臭氧胁迫对大豆氮磷化学计量特征的影响
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摘要
研究臭氧浓度升高对大豆(Glycine max)器官生态化学计量特征的影响,对预测未来生态过程和生物地球化学循环有重要作用。以大豆为试验材料,利用开顶室气室(OTCs)探讨臭氧胁迫对大豆不同器官氮磷化学计量特征的影响。试验设置3个处理:CK处理(对照,O3浓度为45 nmol·mol~(-1))、T1处理(O3浓度为(80±10) nmol·mol~(-1))、T2处理(O_3浓度为(110±10)nmol·mol~(-1))。结果表明,随着生育期的延长,大豆根系和茎秆N含量呈升高趋势,叶片N含量呈先升高后降低的趋势。与对照相比,臭氧浓度升高使根N含量在分枝期和结荚期显著升高(P<0.05),开花期T1处理下显著降低7.13%(P<0.05),T2处理下显著升高17.37%(P<0.05);茎N含量均显著升高,叶片N含量在开花期显著升高,结荚期显著降低。臭氧浓度升高使大豆根、茎、叶P含量表现出不同程度的增加趋势,仅有根系P含量在开花期T1处理下显著降低15.96%(P<0.05),叶片P含量分别在分枝期和结荚期T2处理下显著降低25.10%和11.37%(P<0.05)。茎和根P含量间呈显著正相关(P<0.05),根和茎中N和P含量之间呈显著正相关(P<0.05);N含量和N/P在各器官中表现为叶>根>茎;P含量在各器官中无显著差异。各器官N含量变异系数表现为茎>根>叶;P含量在各器官中变异系数相似;N/P变异系数表现为叶>根>茎。随着臭氧浓度升高和生育期的延长,大豆各器官N/P均小于14,说明大豆生长主要受N素限制。
Studying the effect of elevated ozone concentration on the ecological stoichiometry of soybean(Glycine max) organs plays an important role in predicting future ecological processes and biogeochemical cycles. Open-top chambers(OTCs) were utilized to study the effects on N(nitrogen) and P(phosphorus) stoichiometric characteristics in different organs of soybean under elevated O_3 concentration [(80±10) nmol·mol~(-1) and(110±10) nmol·mol~(-1)]. The results showed that with the extension of growth period, the N content of soybean in roots and stems increased, while the N content in leaves increased firstly and then decreased under enhanced ozone. Compared with the control, elevated ozone resulted in a significant increase in root N content at branching and podding(P<0.05), but a significant decrease of 7.13%(P<0.05) for T1 treatment at flowering and a significant increase of 17.37%(P<0.05) for T2 treatment; N content in stems increased significantly both T1 and T2 treatment; N content in leaves increased at flowering and decreased significantly at podding. On the whole, P content in different organs increased under elevated ozone concentration, but P content in roots decreased significantly 15.96%(P<0.05) at flowering, P content of leaves decreased significantly 25.10% and 11.37%(P<0.05) for T2 treatment both branching and podding respectively. There were significant correlation in P content of root and stem(P<0.05), and N content was significantly correlated with P content in root and stem(P<0.05); N content and N/P ratio of different organs ranked in the order leaf>root>stem, but there was no significant difference in P content among different organs. Meanwhile, the coefficients of variation for N content ranked in the order stem>root>leaf, and the coefficients of variation for N/P ranked in the order root>leaf>stem, but the coefficient of variation for P content was similar in different organs. These results reveal that N/P ratio of soybean organs is less than 14 under ozone stress, indicating that soybean growth is mainly limited by N.
Studying the effect of elevated ozone concentration on the ecological stoichiometry of soybean(Glycine max) organs plays an important role in predicting future ecological processes and biogeochemical cycles. Open-top chambers(OTCs) were utilized to study the effects on N(nitrogen) and P(phosphorus) stoichiometric characteristics in different organs of soybean under elevated O_3 concentration [(80±10) nmol·mol~(-1) and(110±10) nmol·mol~(-1)]. The results showed that with the extension of growth period, the N content of soybean in roots and stems increased, while the N content in leaves increased firstly and then decreased under enhanced ozone. Compared with the control, elevated ozone resulted in a significant increase in root N content at branching and podding(P<0.05), but a significant decrease of 7.13%(P<0.05) for T1 treatment at flowering and a significant increase of 17.37%(P<0.05) for T2 treatment; N content in stems increased significantly both T1 and T2 treatment; N content in leaves increased at flowering and decreased significantly at podding. On the whole, P content in different organs increased under elevated ozone concentration, but P content in roots decreased significantly 15.96%(P<0.05) at flowering, P content of leaves decreased significantly 25.10% and 11.37%(P<0.05) for T2 treatment both branching and podding respectively. There were significant correlation in P content of root and stem(P<0.05), and N content was significantly correlated with P content in root and stem(P<0.05); N content and N/P ratio of different organs ranked in the order leaf>root>stem, but there was no significant difference in P content among different organs. Meanwhile, the coefficients of variation for N content ranked in the order stem>root>leaf, and the coefficients of variation for N/P ranked in the order root>leaf>stem, but the coefficient of variation for P content was similar in different organs. These results reveal that N/P ratio of soybean organs is less than 14 under ozone stress, indicating that soybean growth is mainly limited by N.
引文
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FENG Z Z,HU E Z,WANG X K,et al.,2015.Ground-level O3 pollution and its impacts on food crops in China:a review[J].Environmental Pollution,199(1):42-48.
GUAN L L,WEN D Z,2011.More nitrogen partition in structural proteins and decreased photosynthetic nitrogen-use efficiency of Pinus massoniana under in situ polluted stress[J].Journal of Plant Research,124(6):663-673.
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NIKOLOVA P S,ANDERSEN C P,BLASCHKE H,et al.,2009.Belowground effects of enhanced tropospheric ozone and drought in a beech/spruce forest(Fagus sylvatica L./Picea abies[L.]Karst)[J].Environmental Pollution,158(4):1071-1078.
PiIKKI K,VORNE V,OJANPERA K,et al.,2006.Impact of elevated O3and CO2 exposure on potato(Solanum tuberosum L.cv.Bintje)tuber macronutrients(N,P,K,Mg,Ca)[J].Agriculture Ecosystem and Environment,118(1-4):55-64.
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SHANG B,FENG Z Z,LI P,et al.,2018.Elevated ozone affects C,N and Pecological stoichiometry and nutrient resorption of two poplar clones[J].Environmental Pollution,234:136-144.
SONG Z L,LIU H Y,ZHAO F J,et al.,2014.Ecological stoichiometry of N:P:Si in China’s grasslands[J].Plant and Soil,380(1-2):165-179.
YU H,SHANG H,CHEN Z,TIAN,et al.,2016.Effects of elevated ozone on stoichiometry and nutrient pools of Phoebe Bournei(Hemsl.)Yang and Phoebe Zhennan S.Lee et F.N.Wei seedlings in subtropical China[J].Forests,7(4):78-88.
YUAN Z Y,CHEN H Y,REICH P B,2011.Global-scale latitudinal patterns of plant fine-root nitrogen and phosphorus[J].Nature Communications,2(1):344.
YUAN Z Y,CHEN H Y H,2015.Decoupling of nitrogen and phosphorus in terrestrial plants associated with global changes[J].Nature Climate Change,5(5):465-469.
ZHAO N,YU G R,HE N P,et al.,2016.Invariant allometric scaling of nitrogen and phosphorus in leaves,stems,and fine roots of woody plants along an altitudinal gradient[J].Journal of Plant Research,129(4):647-657.
ZHENG F X,WANG X K,ZHANG W W,et al.,2013.Effects of elevated O3 exposure on nutrient elements and quality of winter wheat and rice grain in Yangtze River Delta,China[J].Environmental Pollution,179(8):19-26.
ZHUANG M H,SHU K L,LI Y C,et al.,2016.Elevated tropospheric ozone affects the concentration and allocation of mineral nutrients of two bamboo species[J].Science of the Total Environment,577:231-235.
鲍士旦,2005.土壤农化分析[M].第3版.北京:中国农业出版社.BAO S D,2005.Soil Agro-chemistrical Analysis[M].The Third Edition.Beijing:China Agriculture Press.
陈娟,曾青,朱建国,等,2011.臭氧和氮肥交互对小麦干物质生产、N、P、K含量及累积量的影响[J].生态环境学报,20(4):616-622.CHEN J,ZENG Q,ZHU J G,et al.,2011 Interactive effects of elevated ozone and nitrogen on dry matter production,concentration and accumulation of nitrogen,phosphorus and potassium in winter wheat[J].Ecology and Environmental Sciences,20(4):616-622.
贺合亮,阳小成,李丹丹,等,2017.青藏高原东部窄叶鲜卑花碳、氮、磷化学计量特征[J].植物生态学报,41(1):126-135.HE H L,YANG X C,LI D D,et al.,2017.Stoichiometric characteristics of carbon,nitrogen and phosphorus of Sibiraea angustata shrub on the eastern Qinghai-Xizang Plateau[J].Chinese Journal of Plant Ecology,41(1):126-135.
贾一磊,2017.臭氧胁迫对不同小麦品种产量、品质和抗倒性的影响[D].扬州:扬州大学:35-41.JIA Y L,2017.Effects of ozone stress on grain yield,quality and plant lodging resistance of different wheat varieties[D].Yangzhou:Yangzhou University:35-41.
刘洋,张健,陈亚梅,等,2013.氮磷添加对巨桉幼苗生物量分配和C?N?P化学计量特征的影响[J].植物生态学报,37(10):933-941.LIU Y,ZHANG J,CHEN Y M,et al.,2013.Effect of nitrogen and phosphorus fertilization on biomass allocation and C?N?Pstoichiometric characteristics of Eucalyptus grandis seedlings[J].Chinese Journal of Plant Ecology,37(10):933-941.
平川,2014.环境变化对兴安落叶松氮磷化学计量特征的影响[D].哈尔滨:东北农业大学:1-44.PING C,2014.Influence of environmental changes on stoichiometric traits of nitrogen and phosphorus for Larix gmelinii trees[D].Harbin:Northeast Forestry University:1-44.
平琴,徐胜,李静,等,2017.坪用白三叶(Trifolium repens)对高浓度臭氧的生理生态响应[J].生态学杂志,36(5):1234-1242.PING Q,XU S,LI J,et al.,2017.Ecophysiological responses of turf-type white clover(Trifolium repens)to elevated O3 concentration[J].Chinese Journal of Ecology,36(5):1234-1242.
孙彩丽,肖列,李鹏,等,2017.氮素添加和干旱胁迫下白羊草碳氮磷化学计量特征[J].植物营养与肥料学报,23(4):1120-1127.SUN C L,XIAO L,LI P,et al.,2017.Effects of nitrogen addition and drought stress on carbon,nitrogen and phosphorus stoichiometry of Bothriochloa ischaemum[J].Journal of Plant Nutrition and Fertilizer.23(4):1120-1127.
王闯,王帅,杨碧波,等,2015.气象条件对沈阳市环境空气臭氧浓度影响研究[J].中国环境监测,31(3):32-37.WANG C,WANG S,YANG B B,et al.,2015.Study of the effect of meteorological conditions on the ambient air ozone concentrations in Shenyang[J].Environmental Monitoring in China,31(3):32-37.
印婧婧,郭大立,何思源,等,2009.内蒙古半干旱区树木非结构性碳、氮、磷的分配格局[J].北京大学学报:自然科学版,45(3):519-527.YIN J J,GUO D L,HE S Y,et al.,2009.Non-structural carbohydrate,N,and P allocation patterns of two temperate tree species in a semi-arid region of Inner Mongolia[J].Acta Scientiarum Naturalium Universitatis Pekinensis,45(3):519-527.
曾德慧,陈广生,2005.生态化学计量学:复杂生命系统奥秘的探索[J].植物生态学报,29(6):1007-1019.ZENG D H,CHEN G S,2005.Ecological stoichiometry:a science to explore the complexity of living systems[J].Acta Phytoecologica Sinica,29(6):1007-1019.
ALLEN,ANDREW P,GILLOOLY,et al.,2009.Towards an integration of ecological stoichiometry and the metabolic theory of ecology to better understand nutrient cycling[J].Ecology Letters,12(5):369-384.
AUSTIN A T,VITOUSEK P M,2012.Introduction to a Virtual Special Issue on ecological stoichiometry and global change[J].The New Phytologist,196(3):649-651.
BYTNEROWICZ A,OMASA K,PAOLETTI E,2007.Integrated effects of air pollution and climate change on forests:a northern hemisphere perspective[J].Environmental Pollution,147(3):438-445.
DUMONT J,KESKISAARI S,KEIN?NEN M,et al.,2014.Ozone affects ascorbate and glutathione biosynthesis as well as amino acid contents in three Euramerican poplar genotypes[J].Tree Physiology,34(3):253-266.
FENG Z Z,HU E Z,WANG X K,et al.,2015.Ground-level O3 pollution and its impacts on food crops in China:a review[J].Environmental Pollution,199(1):42-48.
GUAN L L,WEN D Z,2011.More nitrogen partition in structural proteins and decreased photosynthetic nitrogen-use efficiency of Pinus massoniana under in situ polluted stress[J].Journal of Plant Research,124(6):663-673.
KOERSELMAN W,MEULEMAN A F M,1996.The vegetation N?P ratio:a new tool to detect the nature of nutrient limitation[J].The Journal of Applied Ecology,33(6):1441-1450.
MORGAN P B,BERNACCHI C J,ORT D R,et al.,2004.An in vivo analysis of the effect of season-long open-air elevation of ozone to anticipated 2050 levels on photosynthesis in soybean[J].Plant Physiology,135(4):2348-2357.
NIKOLOVA P S,ANDERSEN C P,BLASCHKE H,et al.,2009.Belowground effects of enhanced tropospheric ozone and drought in a beech/spruce forest(Fagus sylvatica L./Picea abies[L.]Karst)[J].Environmental Pollution,158(4):1071-1078.
PiIKKI K,VORNE V,OJANPERA K,et al.,2006.Impact of elevated O3and CO2 exposure on potato(Solanum tuberosum L.cv.Bintje)tuber macronutrients(N,P,K,Mg,Ca)[J].Agriculture Ecosystem and Environment,118(1-4):55-64.
REICH P B,TJOELKER M G,PREGITZER K S,et al.,2008.Scaling of respiration to nitrogen in leaves,stems and roots of higher land plants[J].Ecology Letters,11(8):793-801.
SHANG B,FENG Z Z,LI P,et al.,2018.Elevated ozone affects C,N and Pecological stoichiometry and nutrient resorption of two poplar clones[J].Environmental Pollution,234:136-144.
SONG Z L,LIU H Y,ZHAO F J,et al.,2014.Ecological stoichiometry of N:P:Si in China’s grasslands[J].Plant and Soil,380(1-2):165-179.
YU H,SHANG H,CHEN Z,TIAN,et al.,2016.Effects of elevated ozone on stoichiometry and nutrient pools of Phoebe Bournei(Hemsl.)Yang and Phoebe Zhennan S.Lee et F.N.Wei seedlings in subtropical China[J].Forests,7(4):78-88.
YUAN Z Y,CHEN H Y,REICH P B,2011.Global-scale latitudinal patterns of plant fine-root nitrogen and phosphorus[J].Nature Communications,2(1):344.
YUAN Z Y,CHEN H Y H,2015.Decoupling of nitrogen and phosphorus in terrestrial plants associated with global changes[J].Nature Climate Change,5(5):465-469.
ZHAO N,YU G R,HE N P,et al.,2016.Invariant allometric scaling of nitrogen and phosphorus in leaves,stems,and fine roots of woody plants along an altitudinal gradient[J].Journal of Plant Research,129(4):647-657.
ZHENG F X,WANG X K,ZHANG W W,et al.,2013.Effects of elevated O3 exposure on nutrient elements and quality of winter wheat and rice grain in Yangtze River Delta,China[J].Environmental Pollution,179(8):19-26.
ZHUANG M H,SHU K L,LI Y C,et al.,2016.Elevated tropospheric ozone affects the concentration and allocation of mineral nutrients of two bamboo species[J].Science of the Total Environment,577:231-235.
鲍士旦,2005.土壤农化分析[M].第3版.北京:中国农业出版社.BAO S D,2005.Soil Agro-chemistrical Analysis[M].The Third Edition.Beijing:China Agriculture Press.
陈娟,曾青,朱建国,等,2011.臭氧和氮肥交互对小麦干物质生产、N、P、K含量及累积量的影响[J].生态环境学报,20(4):616-622.CHEN J,ZENG Q,ZHU J G,et al.,2011 Interactive effects of elevated ozone and nitrogen on dry matter production,concentration and accumulation of nitrogen,phosphorus and potassium in winter wheat[J].Ecology and Environmental Sciences,20(4):616-622.
贺合亮,阳小成,李丹丹,等,2017.青藏高原东部窄叶鲜卑花碳、氮、磷化学计量特征[J].植物生态学报,41(1):126-135.HE H L,YANG X C,LI D D,et al.,2017.Stoichiometric characteristics of carbon,nitrogen and phosphorus of Sibiraea angustata shrub on the eastern Qinghai-Xizang Plateau[J].Chinese Journal of Plant Ecology,41(1):126-135.
贾一磊,2017.臭氧胁迫对不同小麦品种产量、品质和抗倒性的影响[D].扬州:扬州大学:35-41.JIA Y L,2017.Effects of ozone stress on grain yield,quality and plant lodging resistance of different wheat varieties[D].Yangzhou:Yangzhou University:35-41.
刘洋,张健,陈亚梅,等,2013.氮磷添加对巨桉幼苗生物量分配和C?N?P化学计量特征的影响[J].植物生态学报,37(10):933-941.LIU Y,ZHANG J,CHEN Y M,et al.,2013.Effect of nitrogen and phosphorus fertilization on biomass allocation and C?N?Pstoichiometric characteristics of Eucalyptus grandis seedlings[J].Chinese Journal of Plant Ecology,37(10):933-941.
平川,2014.环境变化对兴安落叶松氮磷化学计量特征的影响[D].哈尔滨:东北农业大学:1-44.PING C,2014.Influence of environmental changes on stoichiometric traits of nitrogen and phosphorus for Larix gmelinii trees[D].Harbin:Northeast Forestry University:1-44.
平琴,徐胜,李静,等,2017.坪用白三叶(Trifolium repens)对高浓度臭氧的生理生态响应[J].生态学杂志,36(5):1234-1242.PING Q,XU S,LI J,et al.,2017.Ecophysiological responses of turf-type white clover(Trifolium repens)to elevated O3 concentration[J].Chinese Journal of Ecology,36(5):1234-1242.
孙彩丽,肖列,李鹏,等,2017.氮素添加和干旱胁迫下白羊草碳氮磷化学计量特征[J].植物营养与肥料学报,23(4):1120-1127.SUN C L,XIAO L,LI P,et al.,2017.Effects of nitrogen addition and drought stress on carbon,nitrogen and phosphorus stoichiometry of Bothriochloa ischaemum[J].Journal of Plant Nutrition and Fertilizer.23(4):1120-1127.
王闯,王帅,杨碧波,等,2015.气象条件对沈阳市环境空气臭氧浓度影响研究[J].中国环境监测,31(3):32-37.WANG C,WANG S,YANG B B,et al.,2015.Study of the effect of meteorological conditions on the ambient air ozone concentrations in Shenyang[J].Environmental Monitoring in China,31(3):32-37.
印婧婧,郭大立,何思源,等,2009.内蒙古半干旱区树木非结构性碳、氮、磷的分配格局[J].北京大学学报:自然科学版,45(3):519-527.YIN J J,GUO D L,HE S Y,et al.,2009.Non-structural carbohydrate,N,and P allocation patterns of two temperate tree species in a semi-arid region of Inner Mongolia[J].Acta Scientiarum Naturalium Universitatis Pekinensis,45(3):519-527.
曾德慧,陈广生,2005.生态化学计量学:复杂生命系统奥秘的探索[J].植物生态学报,29(6):1007-1019.ZENG D H,CHEN G S,2005.Ecological stoichiometry:a science to explore the complexity of living systems[J].Acta Phytoecologica Sinica,29(6):1007-1019.
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