氮磷添加对水曲柳化学计量特征和养分再吸收的影响
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摘要
为了解水曲柳林分化学计量特征和养分再吸收率与N、P元素供应的关系,以15年生水曲柳(Fraxinus mandshurica)人工林为研究对象,测定氮(N)、磷(P)添加后水曲柳林分土壤、成熟叶片和衰老叶片养分浓度的变化。结果表明:N、P添加后水曲柳成熟叶N、P浓度和土壤C、N浓度均逐渐增加,而衰老叶P浓度逐渐降低;水曲柳成熟叶C∶N、C∶P和土壤C∶P均随N、P添加而降低,而成熟叶和土壤N∶P仅随P添加而降低,但是衰老叶C∶P、N∶P均随P添加量增加而显著提高; N、P添加后水曲柳叶片N、P再吸收率均提高,其中P再吸收率变化最大,高氮和高磷处理下P再吸收率分别比CK提高2. 3倍和2. 7倍,且叶片P再吸收率与成熟叶C∶P呈显著负相关,但是与衰老叶C∶P、N∶P呈极显著正相关。N、P添加后水曲柳成熟叶N∶P的变化和N、P再吸收率的提高表明:P是限制本研究中水曲柳林木生长的重要因素,N、P添加可以提高水曲柳N、P的利用效率,并改善水曲柳林分的养分状况。
In order to understand the relation between nutrient stoichiometric characteristics and resorption efficiency of Fraxinus mandshurica stand and the N and P supply,N and P concentrations of soils and leaves were determined in the 15-year-old Fraxinus mandshurica plantations after N and P addition. The results showed that: with the increasing of N and P supply,the N and P concentrations of mature leaves and the C and N concentrations of soil increased,while the P concentration of senescent leaves gradually decreased. The C∶ N and C∶ P ratios of mature leaves and C∶ P ratio of soils decreased under N and P addition,and N∶ P ratio of mature leaves and soils decreased under P addition,but C∶ P and N∶ P ratios of senescent leaves increased significantly with the increasing P supply. It showed high resorption efficiency of N and P in leaves under N and P addition,and among them P resorption efficiency were higher. The P resorption efficiency under high nitrogen and high phosphorus treatment increased by 2. 3 times and 2. 7 times of CK,respectively,and the leaf P resorption efficiency and mature leaf C∶ P showed a significantly negative correlation,but it was significantly positively correlated with senescent leaves C∶ P,N∶ P. Our study indicates that N and P addition can adjust N∶ P ratios of mature leaves and increase N and P resorption of Fraxinus mandshurica stand,which suggests that P is an important limiting element of Fraxinus mandshurica during growth,moreover,N and P addition can increase nutrient utilization efficiency and improve nutrient status in the plantation of Fraxinus mandshurica.
In order to understand the relation between nutrient stoichiometric characteristics and resorption efficiency of Fraxinus mandshurica stand and the N and P supply,N and P concentrations of soils and leaves were determined in the 15-year-old Fraxinus mandshurica plantations after N and P addition. The results showed that: with the increasing of N and P supply,the N and P concentrations of mature leaves and the C and N concentrations of soil increased,while the P concentration of senescent leaves gradually decreased. The C∶ N and C∶ P ratios of mature leaves and C∶ P ratio of soils decreased under N and P addition,and N∶ P ratio of mature leaves and soils decreased under P addition,but C∶ P and N∶ P ratios of senescent leaves increased significantly with the increasing P supply. It showed high resorption efficiency of N and P in leaves under N and P addition,and among them P resorption efficiency were higher. The P resorption efficiency under high nitrogen and high phosphorus treatment increased by 2. 3 times and 2. 7 times of CK,respectively,and the leaf P resorption efficiency and mature leaf C∶ P showed a significantly negative correlation,but it was significantly positively correlated with senescent leaves C∶ P,N∶ P. Our study indicates that N and P addition can adjust N∶ P ratios of mature leaves and increase N and P resorption of Fraxinus mandshurica stand,which suggests that P is an important limiting element of Fraxinus mandshurica during growth,moreover,N and P addition can increase nutrient utilization efficiency and improve nutrient status in the plantation of Fraxinus mandshurica.
引文
[1] GUSEWELL S. N∶P ratios in terrestrial plants:variation and functional significance[J]. New Phytologist,2004,164(2):243-266.
[2] BINKLEY D. Forest nutrition management[M]. New York:John Wiley&Sons,Inc,1986.
[3]阎恩荣,王希华,郭明.浙江天童常绿阔叶林、常绿针叶林与落叶阔叶林的C∶N∶P化学计量特征[J].植物生态学报,2010,34(1):48-57.YAN E R,WANG X H,GUO M. C∶N∶P stoichiometry across evergreen broad-leaved forests,evergreen coniferous forests and deciduous broad-leaved forests in the Tiantong region,Zhejiang Province,eastern China[J]. Chinese Journal of Plant Ecology,2010,34(1):48-57.
[4] ZHANG Q F,XIE J S,LYU M K,et al. Short-term effects of soil warming and nitrogen addition on the N∶P stoichiometry of Cunninghamia lanceolata in subtropical regions[J]. Plant&Soil,2017,411:395-407.
[5] KOERSELMAN W,MEULEMAN A F M. The vegetation N∶P ratio:a new tool to detect the nature of nutrient limitation[J]. Journal of Applied Ecology,1996,33(6):1441-1450.
[6] TIAN H Q,CHEN G S,ZHANG C,et al. Pattern and variation of C∶N∶P ratios in China’s soils:A synthesis of observational data[J]. Biogeochemistry,2010,98(3):139-151.
[7]黄菊莹,余海龙.不同氮磷比处理对甘草生长与生态化学计量特征的影响[J].植物生态学报,2017,41(3):325-336.HUANG J Y,YU H L. Effects of different nitrogen:phosphorus levels on the growth and ecological stoichiometry of Glycyrrhiza uralensis[J]. Chinese Journal of Plant Ecology,2017,41(3):325-336.
[8]刘芳,葛江丽,施汉钰,等.西洋参地氮磷钾含量变化规律研究[J].林业科技,2017,42(6):19-21.LIU F,GE J L,SHI H Y,et al. Study on changing rule of nitrogen,phosphorus and potassium content in Panax quinquefolium field[J]. Forestry Science&Technology,2017,42(6):19-21.
[9] TESSIER J T,RAYNAL D J. Use of nitrogen to phosphorus ratios in plant tissue as an indicator of nutrient limitation and nitrogen saturation[J]. Journal of Applied Ecology,2003,40(3):523-534.
[10] ELSER J J,DOBBERFUHL D R,MACKAY N A,et al. Organism size,life history,and N:P stoichiometry[J]. Bio Science,1996,46(9):674-684.
[11] YOU C M,WU F Z,YANG W Q,et al. Nutrient-limited conditions determine the responses of foliar nitrogen and phosphorus stoichiometry to nitrogen addition:A global meta-analysis[J].Environmental Pollution,2018,241:740-749.
[12] MO Q F,ZOU B,LI Y W,et al. Response of plant nutrient stoichiometry to fertilization varied with plant tissues in a tropical forest[J]. Scientific Reports,2015,5(1):14605.
[13] KILLINGBECK K T. The terminological jungle revisited:Making a case for use of the term resorption[J]. Oikos,1986,46(2):263-264.
[14] AERTS R. Nutrient use efficiency in evergreen and deciduous species[J]. Oecologia,1990,84(3):391-397.
[15] REED S C,TOWNSEND A R,DAVIDSON E A,et al. Stoichiometric patterns in foliar nutrient resorption across multiple scales[J]. New Phytologist,2012,196(1):173-180.
[16]赵琼,刘兴宇,胡亚林.氮添加对兴安落叶松养分分配和再吸收效率的影响[J].林业科学,2010,46(5):14-19.ZHAO Q,LIU X Y,HU Y L. Effects of nitrogen addition on nutrient allocation and nutrient resorption efficiency in Larix gmelinii[J]. Scientia Silvae Sinicae,2010,46(5):14-19.
[17] SON Y W,LEE I K,RYU S R. Nitrogen and phosphorus dynamics in foliage and twig of pitch pine and Japanese larch plantations in relation to fertilization[J]. Journal of Plant Nutrition,2000,23(5):697-710.
[18] CHAPIN F S,MIOLANEN L. Nutritional controls over nitrogen and phosphorus resorption from Alaskan birch leaves[J]. Ecology,1991,72(2):709-715.
[19] VAN HEERWAARDEN L M,TOET S,AERTS R. Nitrogen and phosphorus resorption efficiency and proficiency in six sub-arctic bog species after 4 years of nitrogen fertilization[J]. Journal of Ecology,2003,91(6):1060-1070.
[20] ZHANG L X,BAI Y F. Differential responses of N:P stoichiometry of Leymus chinensis and Carex korshinskyi to N additions in a steppe ecosystem in Nei Mongol[J]. Acta Botanica Sinica,2004,46(3):259-270.
[21]范志强,沈海龙,王庆成,等.水曲柳幼林适生立地条件研究[J].林业科学,2002,38(2):38-43.FAN Z Q,SHEN H L,WANG Q C,et al. Study on the conditions suitable for young plantation of Fraxinus mandshurica[J].Scientia Silvae Sinicae,2002,38(2):38-43.
[22]那萌,刘婷岩,张彦东,等.林分密度对水曲柳人工林碳储量的影响[J].北京林业大学学报,2017,39(1):20-26.NA M,LIU T Y,ZHANG Y D,et al. Effects of stock density on carbon storage in Fraxinus mandshurica plantations[J]. Journal of Beijing Forestry University,2017,39(1):20-26.
[23]郝玉琢,王树力.水曲柳人工纯林与混交林土壤生态化学计量特征的比较[J].东北林业大学学报,2018,46(3):54-58.HAO Y Z,WANG S L. Soil ecological stoichiometric characteristics between pure Fraxinus mandshurica plantation and mixed F.mandshurica plantation[J]. Journal of Northeast Forestry University,2018,46(3):54-58.
[24]陆姣云,段兵红,杨梅,等.植物叶片氮磷养分重吸收规律及其调控机制研究进展[J].草业学报,2018,27(4):178-188.LU J Y,DUAN B H,YANG M,et al. Research progress in nitrogen and phosphorus resorption from senesced leaves and the influence of ontogenetic and environmental factors[J]. Acta Prataculturae Sinica,2018,27(4):178-188.
[25] SPARKS D L,PAGE A L,LOEPPERT P A,et al. Methods of soil analysis,part 3:chemical methods[M]. Madison:Soil Science Society of America and American Society of Agronomy,1996.
[26] KILLINGBECK K T. Nutrients in senesced leaves:keys to the search for potential resorption and resorption proficiency[J]. Ecology,1996,77(6):1716-1727.
[27]王淑平,周广胜,吕育财.中国东北样带(NECT)土壤碳,氮,磷的梯度分布及其与气候因子的关系[J].植物生态学报,2002,26(5):513-517.WANG S P,ZHOU G S,LV Y C. Distribution of soil carbon,nitrogen and phosphorus along northeast China transect(NECT)and their relationships with climatic factors[J]. Acta Phytoecologica Sinica,2002,26(5):513-517.
[28]陈伏生,胡小飞,葛刚.城市地被植物麦冬叶片氮磷化学计量比和养分再吸收效率[J].草业学报,2007,16(4):47-54.CHEN F S,HU X F,GE G. Nitrogen and phosphorus stoichiometry and nutrient resorption efficiency of urban ground cover[J].Acta Prataculturae Sinica,2007,16(4):47-54.
[29]曹昊,刘瑰琦,施汉钰,等.氮、磷、钾肥施肥量对半夏产量影响的研究[J].林业科技,2017,42(2):35-37.CAO H,LIU G Q,SHI H Y,et al. Effect of nitrogen,phosphorus and potassium fertilizer application rate on pinellia ternata yield[J]. Forestry Science&Technology,2017,42(2):35-37.
[30] CORREIA P J,MARTINSLOUCAO M A. Leaf nutrient variation in mature carob(Ceratonia siliqua)trees in response to irrigation and fertilization[J]. Tree Physiology,1997,17(12):813-819.
[31] WEI H X,XU C Y,MA L Y,et al. Short-term nitrogen(N)-retranslocation within Larix olgensis seedlings is driven to increase by N-deposition:evidence from a simulated 15N experiment in northeast China[J]. International Journal of Agriculture&Biology,2014,16(6):1031-1040.
[32]邓浩俊,陈爱民,严思维,等.不同林龄新银合欢重吸收率及其C∶N∶P化学计量特征[J].应用与环境生物学报,2015,21(3):522-527.DENG H J,CHEN A M,YAN S W,et al. Nutrient resorption efficiency and C∶N∶P stoichiometry in different ages of Leucaena leucocephala[J]. Chinese Journal of Applied Environmental Biology,2015,21(3):522-527.
[2] BINKLEY D. Forest nutrition management[M]. New York:John Wiley&Sons,Inc,1986.
[3]阎恩荣,王希华,郭明.浙江天童常绿阔叶林、常绿针叶林与落叶阔叶林的C∶N∶P化学计量特征[J].植物生态学报,2010,34(1):48-57.YAN E R,WANG X H,GUO M. C∶N∶P stoichiometry across evergreen broad-leaved forests,evergreen coniferous forests and deciduous broad-leaved forests in the Tiantong region,Zhejiang Province,eastern China[J]. Chinese Journal of Plant Ecology,2010,34(1):48-57.
[4] ZHANG Q F,XIE J S,LYU M K,et al. Short-term effects of soil warming and nitrogen addition on the N∶P stoichiometry of Cunninghamia lanceolata in subtropical regions[J]. Plant&Soil,2017,411:395-407.
[5] KOERSELMAN W,MEULEMAN A F M. The vegetation N∶P ratio:a new tool to detect the nature of nutrient limitation[J]. Journal of Applied Ecology,1996,33(6):1441-1450.
[6] TIAN H Q,CHEN G S,ZHANG C,et al. Pattern and variation of C∶N∶P ratios in China’s soils:A synthesis of observational data[J]. Biogeochemistry,2010,98(3):139-151.
[7]黄菊莹,余海龙.不同氮磷比处理对甘草生长与生态化学计量特征的影响[J].植物生态学报,2017,41(3):325-336.HUANG J Y,YU H L. Effects of different nitrogen:phosphorus levels on the growth and ecological stoichiometry of Glycyrrhiza uralensis[J]. Chinese Journal of Plant Ecology,2017,41(3):325-336.
[8]刘芳,葛江丽,施汉钰,等.西洋参地氮磷钾含量变化规律研究[J].林业科技,2017,42(6):19-21.LIU F,GE J L,SHI H Y,et al. Study on changing rule of nitrogen,phosphorus and potassium content in Panax quinquefolium field[J]. Forestry Science&Technology,2017,42(6):19-21.
[9] TESSIER J T,RAYNAL D J. Use of nitrogen to phosphorus ratios in plant tissue as an indicator of nutrient limitation and nitrogen saturation[J]. Journal of Applied Ecology,2003,40(3):523-534.
[10] ELSER J J,DOBBERFUHL D R,MACKAY N A,et al. Organism size,life history,and N:P stoichiometry[J]. Bio Science,1996,46(9):674-684.
[11] YOU C M,WU F Z,YANG W Q,et al. Nutrient-limited conditions determine the responses of foliar nitrogen and phosphorus stoichiometry to nitrogen addition:A global meta-analysis[J].Environmental Pollution,2018,241:740-749.
[12] MO Q F,ZOU B,LI Y W,et al. Response of plant nutrient stoichiometry to fertilization varied with plant tissues in a tropical forest[J]. Scientific Reports,2015,5(1):14605.
[13] KILLINGBECK K T. The terminological jungle revisited:Making a case for use of the term resorption[J]. Oikos,1986,46(2):263-264.
[14] AERTS R. Nutrient use efficiency in evergreen and deciduous species[J]. Oecologia,1990,84(3):391-397.
[15] REED S C,TOWNSEND A R,DAVIDSON E A,et al. Stoichiometric patterns in foliar nutrient resorption across multiple scales[J]. New Phytologist,2012,196(1):173-180.
[16]赵琼,刘兴宇,胡亚林.氮添加对兴安落叶松养分分配和再吸收效率的影响[J].林业科学,2010,46(5):14-19.ZHAO Q,LIU X Y,HU Y L. Effects of nitrogen addition on nutrient allocation and nutrient resorption efficiency in Larix gmelinii[J]. Scientia Silvae Sinicae,2010,46(5):14-19.
[17] SON Y W,LEE I K,RYU S R. Nitrogen and phosphorus dynamics in foliage and twig of pitch pine and Japanese larch plantations in relation to fertilization[J]. Journal of Plant Nutrition,2000,23(5):697-710.
[18] CHAPIN F S,MIOLANEN L. Nutritional controls over nitrogen and phosphorus resorption from Alaskan birch leaves[J]. Ecology,1991,72(2):709-715.
[19] VAN HEERWAARDEN L M,TOET S,AERTS R. Nitrogen and phosphorus resorption efficiency and proficiency in six sub-arctic bog species after 4 years of nitrogen fertilization[J]. Journal of Ecology,2003,91(6):1060-1070.
[20] ZHANG L X,BAI Y F. Differential responses of N:P stoichiometry of Leymus chinensis and Carex korshinskyi to N additions in a steppe ecosystem in Nei Mongol[J]. Acta Botanica Sinica,2004,46(3):259-270.
[21]范志强,沈海龙,王庆成,等.水曲柳幼林适生立地条件研究[J].林业科学,2002,38(2):38-43.FAN Z Q,SHEN H L,WANG Q C,et al. Study on the conditions suitable for young plantation of Fraxinus mandshurica[J].Scientia Silvae Sinicae,2002,38(2):38-43.
[22]那萌,刘婷岩,张彦东,等.林分密度对水曲柳人工林碳储量的影响[J].北京林业大学学报,2017,39(1):20-26.NA M,LIU T Y,ZHANG Y D,et al. Effects of stock density on carbon storage in Fraxinus mandshurica plantations[J]. Journal of Beijing Forestry University,2017,39(1):20-26.
[23]郝玉琢,王树力.水曲柳人工纯林与混交林土壤生态化学计量特征的比较[J].东北林业大学学报,2018,46(3):54-58.HAO Y Z,WANG S L. Soil ecological stoichiometric characteristics between pure Fraxinus mandshurica plantation and mixed F.mandshurica plantation[J]. Journal of Northeast Forestry University,2018,46(3):54-58.
[24]陆姣云,段兵红,杨梅,等.植物叶片氮磷养分重吸收规律及其调控机制研究进展[J].草业学报,2018,27(4):178-188.LU J Y,DUAN B H,YANG M,et al. Research progress in nitrogen and phosphorus resorption from senesced leaves and the influence of ontogenetic and environmental factors[J]. Acta Prataculturae Sinica,2018,27(4):178-188.
[25] SPARKS D L,PAGE A L,LOEPPERT P A,et al. Methods of soil analysis,part 3:chemical methods[M]. Madison:Soil Science Society of America and American Society of Agronomy,1996.
[26] KILLINGBECK K T. Nutrients in senesced leaves:keys to the search for potential resorption and resorption proficiency[J]. Ecology,1996,77(6):1716-1727.
[27]王淑平,周广胜,吕育财.中国东北样带(NECT)土壤碳,氮,磷的梯度分布及其与气候因子的关系[J].植物生态学报,2002,26(5):513-517.WANG S P,ZHOU G S,LV Y C. Distribution of soil carbon,nitrogen and phosphorus along northeast China transect(NECT)and their relationships with climatic factors[J]. Acta Phytoecologica Sinica,2002,26(5):513-517.
[28]陈伏生,胡小飞,葛刚.城市地被植物麦冬叶片氮磷化学计量比和养分再吸收效率[J].草业学报,2007,16(4):47-54.CHEN F S,HU X F,GE G. Nitrogen and phosphorus stoichiometry and nutrient resorption efficiency of urban ground cover[J].Acta Prataculturae Sinica,2007,16(4):47-54.
[29]曹昊,刘瑰琦,施汉钰,等.氮、磷、钾肥施肥量对半夏产量影响的研究[J].林业科技,2017,42(2):35-37.CAO H,LIU G Q,SHI H Y,et al. Effect of nitrogen,phosphorus and potassium fertilizer application rate on pinellia ternata yield[J]. Forestry Science&Technology,2017,42(2):35-37.
[30] CORREIA P J,MARTINSLOUCAO M A. Leaf nutrient variation in mature carob(Ceratonia siliqua)trees in response to irrigation and fertilization[J]. Tree Physiology,1997,17(12):813-819.
[31] WEI H X,XU C Y,MA L Y,et al. Short-term nitrogen(N)-retranslocation within Larix olgensis seedlings is driven to increase by N-deposition:evidence from a simulated 15N experiment in northeast China[J]. International Journal of Agriculture&Biology,2014,16(6):1031-1040.
[32]邓浩俊,陈爱民,严思维,等.不同林龄新银合欢重吸收率及其C∶N∶P化学计量特征[J].应用与环境生物学报,2015,21(3):522-527.DENG H J,CHEN A M,YAN S W,et al. Nutrient resorption efficiency and C∶N∶P stoichiometry in different ages of Leucaena leucocephala[J]. Chinese Journal of Applied Environmental Biology,2015,21(3):522-527.