荒漠黑果枸杞化学计量特征对氮磷添加比例和水平的响应

doi:10.12118/j.issn.1000–6060.2021.06.23

摘要/Abstract

摘要：

植物化学计量特征对阐明生物地球化学及生态过程对全球变化的响应至关重要。研究不同氮磷（N、P）添加处理对荒漠植物生态化学计量特征的影响,可从化学计量的角度理解植物对环境变化的响应,为预测全球变化背景下植物和营养元素的相互作用提供思路。以黑果枸杞（Lycium ruthenicum）为材料,设置3个氮磷添加量和比例,通过大田试验研究了黑果枸杞C:N:P化学计量特征对氮磷添加的响应,比较分析了器官间的化学计量特征。结果表明：（1）氮磷添加对黑果枸杞不同器官碳（C）含量的影响较小,随氮磷添加量的增加,细根N含量显著增大;随氮磷添加比例的提高,非根器官N含量显著增加,根系P含量显著降低;氮磷添加比例和添加量的交互效应显著影响了根系和果实N、P含量及茎P含量。（2）低氮磷添加比处理降低了器官碳磷比（C/P）,提高了碳氮比（C/N）,而高氮磷添加比处理与之相反,各器官氮磷比（N/P）维持在相对稳定的水平,黑果枸杞通过调整养分保存策略以保守的养分利用方式抵消环境元素化学计量的变化。（3）黑果枸杞化学计量特征体现出器官功能差异性,叶片N含量和N/P显著高于其他器官,茎C/N最高,粗根C含量和C/P最高,细根N、P含量和N/P较高,果实P含量最高。相对于C,代谢活跃器官（叶、细根、果实）比非代谢活跃器官（茎、粗根）需要更多的N和P。研究结果有助于更好地从元素与植物功能的角度理解荒漠植物化学计量特征对氮磷添加的响应。

关键词: 化学计量特征, 氮磷添加量, 氮磷添加比例, 黑果枸杞

Abstract:

The stoichiometric features of plants are important for elucidating the response of biogeochemical and ecological methods to global change. Studying the effects of different nitrogen (N) and phosphorus (P) addition treatments on the ecological stoichiometric characteristics of desert plants can help understand the response of plants to environmental changes from the perspective of stoichiometry and provide ideas for predicting plants and nutrients interactions in the context of global change. Therefore, we used Lycium ruthenicum as the material and set up three N, P supply ratios and supply levels and examined the effects of the supply ratio and level on the C:N:P stoichiometric features of various organs. The results indicated that: (1) Nitrogen and phosphorus addition had little impact on the carbon (C) content in various organs. The content of N in the fine roots considerably increased with a rise in N, P supply level. With the increase in N, P supply ratio, the content of N in nonroot organs increased and the content of P in the root system decreased considerably. The interaction between the proportion of N, P supply level and the supply ratio greatly affected the N, P content in roots and fruits as well as the P content in stems. (2) The low N/P treatment decreased organ C/P and increased C/N, whereas the high N/P treatment was the opposite. The N/P of each organ was maintained at a relatively stable level, Lycium ruthenicum offset the change in environmental element stoichiometry in a conservative nutrient utilization method by altering the nutrient preservation strategy. (3) Stoichiometric characteristics reflected organ differences. The content of N and N/P in leaves was considerably higher than that in other organs. The content of C/N in stems, C and P in coarse roots, as well as the content of P in fruits was the highest. The NP and N/P contents in fine roots were also higher. Compared with C, metabolically active organs (leaves, fine roots, fruits) require more N and P than structural organs (stems, coarse roots). From the perspective of elements and plant functions, the research findings were helpful in better understanding the response of desert plant stoichiometric properties to nitrogen and phosphorus addition.

Key words: stoichiometry characteristics, N, P supply level, N, P supply ratio, Lycium ruthenicum

李金霞,陈年来,孙小妹,李发奎,郁洁汝. 荒漠黑果枸杞化学计量特征对氮磷添加比例和水平的响应[J]. 干旱区地理, 2021, 44(6): 1750-1762.

LI Jinxia,CHEN Nianlai,SUN Xiaomei,LI Fakui,YU Jieru. Response of stoichiometry of desert Lycium ruthenicum to N and P addition levels and ratios[J]. Arid Land Geography, 2021, 44(6): 1750-1762.

图/表 9

表1

图1

图2

图3

图4

图5

表2

图6

图7

参考文献 51

[1]	周欣, 左小安, 赵学勇, 等. 科尔沁沙地不同生境植物及叶片的C、N元素计量特征[J]. 干旱区地理, 2015, 38(3):565-575.
	[ Zhou Xin, Zuo Xiao’an, Zhao Xueyong, et al. Ecological stoichiometry of plant and leaf carbon and nitrogen in different habitats of Horqin Sandy Land[J]. Arid Land Geography, 2015, 38(3):565-575. ]
[2]	Sterner R W, Elser J J. Ecological stoichiometry: The biology of elements from molecules to the biosphere[M]. Princeton: Princeton University Press, 2002.
[3]	李从娟, 徐新文, 孙永强, 等. 不同生境下三种荒漠植物叶片及土壤C、N、P的化学计量特征[J]. 干旱区地理, 2014, 37(5):996-1004.
	[ Li Congjuan, Xu Xinwen, Sun Yongqiang, et al. Stoichiometric characteristics of C, N, P for three desert plants leaf and soil at different habitats[J]. Arid Land Geography, 2014, 37(5):996-1004. ]
[4]	Peñuelas J, Poulter B, Sardans J, et al. Human-induced nitrogen-phosphorus imbalances alter natural and managed ecosystems across the globe[J]. Nature Communications, 2013, 4(1):2934, doi: 10.1038/ncomms3934. doi: 10.1038/ncomms3934
[5]	Imran M, Gurmani Z A. Role of macro and micro nutrients in the plant growth and development[J]. Science, Technology and Development, 2011, 30(3):36-40.
[6]	Marklein A R, Houlton B Z. Nitrogen inputs accelerate phosphorus cycling rates across a wide variety of terrestrial ecosystems[J]. New Phytologist, 2012, 193(3):696-704. doi: 10.1111/j.1469-8137.2011.03967.x pmid: 22122515
[7]	Huang W J, Houlton B Z, Marklein A R, et al. Plant stoichiometric responses to elevated CO₂ vary with nitrogen and phosphorus inputs: Evidence from a global-scale meta-analysis[J]. Scientific Reports, 2015, 5(1):1-8.
[8]	Liu J X, Fang X, Deng Q, et al. CO₂ enrichment and N addition increase nutrient loss from decomposing leaf litter in subtropical model forest ecosystems[J]. Scientific Reports, 2015(5):7952, doi: 10.1038/srep07952. doi: 10.1038/srep07952
[9]	Zhang S B, Zhang J L Slik J W F, et al. Leaf element concentrations of terrestrial plants across China are influenced by taxonomy and the environment[J]. Global Ecology & Biogeography, 2012, 21(8):809-818.
[10]	Sardans J, Rivas-Ubach A, Peñuelas J. The C:N:P stoichiometry of organisms and ecosystems in a changing world: A review and perspectives[J]. Perspectives in Plant Ecology Evolution & Systematics, 2012, 14(1):33-47.
[11]	孙小妹, 何明珠, 周彬, 等. 霸王根茎叶非结构性碳与C:N:P计量特征对干旱的响应[J]. 干旱区地理, 2021, 44(1):240-249.
	[ Sun Xiaomei, He Mingzhu, Zhou Bin, et al. Non-structural carbohydrates and C:N:P stoichiometry of roots, stems, and leaves of Zygophyllum xanthoxylon in responses to xeric condition[J]. Arid Land Geography, 2021, 44(1):240-249. ]
[12]	Zhao W Q, Reich P B, Yu Q N, et al. Shrub type dominates the vertical distribution of leaf C:N:P stoichiometry across an extensive altitudinal gradient[J]. Biogeoences, 2018, 15:2033-2053.
[13]	Kerkhoff A J, Fagan W F, Elser J J, et al. Phylogenetic and growth form variation in the scaling of nitrogen and phosphorus in the seed plants[J]. The American Naturalist, 2006, 168(4):E103-22, doi: 10.1086/507879. doi: 10.1086/507879
[14]	Agren G. Stoichiometry and nutrition of plant growth in natural communities[J]. Annual Review of Ecology Evolution & Systematics, 2008, 39(1):153-170.
[15]	Minden V, Kleyer M. Internal and external regulation of plant organ stoichiometry[J]. Plant Biology, 2014, 16(5):897-907. doi: 10.1111/plb.12155 pmid: 24552639
[16]	Güsewell S. N:P ratios in terrestrial plants: Variation and functional significance[J]. New Phytologist, 2010, 164(2):243-266. doi: 10.1111/nph.2004.164.issue-2
[17]	Xu Z H, Chen C R, He J Z, et al. Trends and challenges in soil research 2009: Linking global climate change to local long-term forest productivity[J]. Journal of Soils and Sediments, 2009, 9(2):83-88. doi: 10.1007/s11368-009-0060-6
[18]	Elser J J, Fagan W F, Kerkhoff A J, et al. Biological stoichiometry of plant production: Metabolism, scaling and ecological response to global change[J]. New Phytologist, 2010, 186(3):593-608. doi: 10.1111/j.1469-8137.2010.03214.x pmid: 20298486
[19]	王春成, 马松梅, 张丹, 等. 柴达木野生黑果枸杞的空间遗传结构[J]. 植物生态学报, 2020, 44(6):661-668. doi: 10.17521/cjpe.2019.0298
	[ Wang Chuncheng, Ma Songmei, Zhang Dan, et al. Spatial genetic structure of Lycium ruthenicum in the Qaidam Basin[J]. Chinese Journal of Plant Ecology, 2020, 44(6):661-668. ] doi: 10.17521/cjpe.2019.0298
[20]	Güsewell S. High nitrogen: Phosphorus ratios reduce nutrient retention and second-year growth of wetland sedges[J]. New Phytologist, 2005, 166(2):537-550. pmid: 15819916
[21]	鲍士旦. 土壤农化分析[M]. 第3版. 北京: 中国农业出版社, 2007: 268-270, 389-391.
	[ Bao Shidan. Soil agro-chemistrical analysis[M]. 3rd ed. Beijing: China Agriculture Press, 2007: 268-270, 389-391. ]
[22]	Aerts R, Chapin F. The mineral nutrition of wild plants revisited: A re-evaluation of processes and patterns[J]. Advances in Ecological Research, 1999, 30:1-67.
[23]	Jing H, Zhou H X, Wang G L, et al. Nitrogen addition changes the stoichiometry and growth rate of different organs in Pinus tabuliformis seedlings[J]. Frontiers in Plant Science. 2017, 8:1922, doi: 10.3389/fpls.2017.01922. doi: 10.3389/fpls.2017.01922
[24]	Cui Q, Xiao T L, Wang Q B, et al. Nitrogen fertilization and fire act independently on foliar stoichiometry in a temperate steppe[J]. Plant & Soil, 2010, 334(1):209-219.
[25]	石贤萌, 杞金华, 宋亮, 等. 哀牢山中山湿性常绿阔叶林两种优势幼苗C、N、P化学计量特征及其对N沉降增加的响应[J]. 植物生态学报, 2015, 39(10):962-970. doi: 10.17521/cjpe.2015.0093
	[ Shi Xianmeng, Qi Jinhua, Song liang, et al. C, N and P stoichiometry of two dominant seedlings and their responses to nitrogen additions in the montane moist evergreen broad-leaved forest in Ailao Mountains, Yunnan[J]. Chinese Journal of Plant Ecology, 2015, 39(10):962-970. ] doi: 10.17521/cjpe.2015.0093
[26]	余华, 潘宗涛, 陈志强, 等. 氮添加对侵蚀退化红壤化学性质及芒萁叶功能性状的影响[J]. 应用与环境生物学报, 2020, 26(4):1-11.
	[ Yu Hua, Pan Zongtao, Chen Zhiqiang, et al. Effects of nitrogen addition on soil chemical properties and leaf functional traits of Dicranopteris dichotoma in the red soil erosion area of southern China[J]. Chinese Journal of Applied and Environmental Biology, 2020, 26(4):1-11. ]
[27]	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, 5:14605, doi: 10.1038/srep14605. doi: 10.1038/srep14605
[28]	Elser J J, Fagan W F, Denno R F, et al. Nutritional constraints in terrestrial and freshwater food webs[J]. Nature, 2000, 408(6812):578-580. doi: 10.1038/35046058
[29]	Sistla S A, Schimel J P. Stoichiometric flexibility as a regulator of carbon and nutrient cycling in terrestrial ecosystems under change[J]. New Phytologist, 2012, 196(1):68-78. doi: 10.1111/j.1469-8137.2012.04234.x pmid: 22924404
[30]	Sardans J, Peñuelas J. Climate and taxonomy underlie different elemental concentrations and stoichiometries of forest species: The optimum “biogeochemical niche”[J]. Plant Ecology, 2014, 215(4):441-455. pmid: 25983614
[31]	Persson J, Fink P, Goto A, et al. To be or not to be what you eat: Regulation of stoichiometric homeostasis among autotrophs and heterotrophs[J]. Oikos, 2010, 119(5):741-751. doi: 10.1111/j.1600-0706.2009.18545.x
[32]	Huang J Y, Wang P, Niu Y B, et al. Changes in C:N:P stoichiometry modify N and P conservation strategies of a desert steppe species Glycyrrhiza uralensis[J]. Scientific Reports, 2018, 8(1):1-9.
[33]	Griffiths B S, Spilles A, Bonkowski M. C:N:P stoichiometry and nutrient limitation of the soil microbial biomass in a grazed grassland site under experimental P limitation or excess[J]. Ecological Processes, 2012, 1:6, doi: 10.1186/2192-1709-1-6. doi: 10.1186/2192-1709-1-6
[34]	Wang Z N, Lu J Y, Yang M, et al. Stoichiometric characteristics of carbon, nitrogen, and phosphorus in leaves of differently aged lucerne (Medicago sativa) stands[J]. Frontiers in Plant Science, 2015, 6:1062, doi: 10.3389/fpls.2015.01062. doi: 10.3389/fpls.2015.01062
[35]	Tian H, Chen G, Zhang C, et al. Pattern and variation of C:N:P ratios in China’s soils: A synjournal of observational data[J]. Biogeochemistry, 2010, 98(1/3):139-151. doi: 10.1007/s10533-009-9382-0
[36]	梁星云. 长白山阔叶红松林演替系列主要树种叶片功能性状与化学计量学研究[D]. 北京: 中国林业科学研究院, 2017.
	[ Liang Xingyun. Leaf functional traits and ecological stoichiometry of the dominant tree species along forest succession of Korean pine and broad-leaved mixed forest in Changbai Mountain, northeastern China[D]. Beijing: Chinese Academy of Forestry, 2017. ]
[37]	Li W B, Jin C J, Guan D X, et al. The effects of simulated nitrogen deposition on plant root traits: A meta-analysis[J]. Soil Biology & Biochemistry, 2015, 82:112-118. doi: 10.1016/j.soilbio.2015.01.001
[38]	Li L, Liu B, Gao X P, et al. Nitrogen and phosphorus addition differentially affect plant ecological stoichiometry in desert grassland[J]. Scientific Reports, 2019, 9:18673, doi: 10.1038/s41598-019-55275-8. doi: 10.1038/s41598-019-55275-8
[39]	Zhan S X, Wang Y, Zhu Z C, et al. Nitrogen enrichment alters plant N:P stoichiometry and intensifies phosphorus limitation in a steppe ecosystem[J]. Environmental and Experimental Botany, 2017, 134:21-32. doi: 10.1016/j.envexpbot.2016.10.014
[40]	Chen G T, Tu L H, Peng Y, et al. Effect of nitrogen additions on root morphology and chemistry in a subtropical bamboo forest[J]. Plant Soil, 2017, 412:441-451. doi: 10.1007/s11104-016-3074-z
[41]	Mayor J R, Wright S J, Turner B L. Species-specific responses of foliar nutrients to long-term nitrogen and phosphorus additions in a lowland tropical forest[J]. Journal of Ecology, 2014, 102(1):36-44. doi: 10.1111/jec.2013.102.issue-1
[42]	Kleyer M, Minden V. Why functional ecology should consider all plant organs: An allocation-based perspective[J]. Basic and Applied Ecology, 2014, 16(1):1-9. doi: 10.1016/j.baae.2014.11.002
[43]	Reich P B, Oleksyn J. Global patterns of plant leaf N and P in relation to temperature and latitude[J]. Proceedings of the National Academy of Sciences of the United States of America, 2004, 101(30):11001-11006.
[44]	Han W X, Fang J Y, Guo D L, et al. Leaf nitrogen and phosphorus stoichiometry across 753 terrestrial plant species in China[J]. New Phytologist, 2005, 168(2):377-385. doi: 10.1111/nph.2005.168.issue-2
[45]	李玉霖, 毛伟, 赵学勇, 等. 北方典型荒漠及荒漠化地区植物叶片氮磷化学计量特征研究[J]. 环境科学, 2010, 31(8):1716-1725.
	[ Li Yulin, Mao Wei, Zhao Xueyong, et al. Leaf nitrogen and phosphorus stoichiometry in typical desert and desertified regions, north China[J]. Environmental Science, 2010, 31(8):1716-1725. ]
[46]	Li H, Li J, He Y L, et al. Changes in carbon, nutrients and stoichiometric relations under different soil depths, plant tissues and ages in black locust plantations[J]. Acta Physiologiae Plantarum, 2013, 35(10):2951-2964. doi: 10.1007/s11738-013-1326-6
[47]	马玉珠, 钟全林, 靳冰洁, 等. 中国植物细根碳、氮、磷化学计量学的空间变化及其影响因子[J]. 植物生态学报, 2015, 39(2):159-166. doi: 10.17521/cjpe.2015.0015
	[ Ma Yuzhu, Zhong Quanlin, Jin Bingjie, et al. Spatial changes and influencing factors of fine root carbon, nitrogen and phosphorus stoichiometry of plants in China[J]. Chinese Journal of Plant Ecology, 2015, 39(2):159-166. ] doi: 10.17521/cjpe.2015.0015
[48]	Freschet G T, Cornwell W K, Wardle D, et al. Linking litter decomposition of above-and below-ground organs to plant-soil feedbacks worldwide[J]. Journal of Ecology, 2013, 101(4):943-952. doi: 10.1111/1365-2745.12092
[49]	Wang G, Fahey T J, Xue S, et al. Root morphology and architecture respond to N addition in Pinus tabuliformis, west China[J]. Oecologia, 2012, 171(2):583-590. doi: 10.1007/s00442-012-2441-6
[50]	Wang G L, Liu F. Carbon allocation of Chinese pine seedlings along a nitrogen addition gradient[J]. Forest Ecology and Management, 2014, 334:114-121. doi: 10.1016/j.foreco.2014.09.004
[51]	Treseder K K, Vitousek P M. Effects of soil nutrient availability on investment in acquisition of N and P in Hawaiian rain forests[J]. Ecology, 2001, 82(4):946-954. doi: 10.1890/0012-9658(2001)082[0946:EOSNAO]2.0.CO;2

器官		碳（C）	氮（N）	磷（P）	碳氮比（C/N）	碳磷比（C/P）	氮磷比（N/P）
粗根	添加比（SR）	6.057^*	2.409	14.701^***	3.254	11.032^**	13.070^***
	添加量（SL）	2.675	3.991^*	12.135^***	4.433^*	9.045^**	3.467
	SR×SL	3.033^*	3.014^*	5.595^**	1.743	6.060^**	6.381^**
细根	添加比（SR）	2.948	0.134	38.262^***	0.968	43.815^***	39.461^***
	添加量（SL）	1.804	1.660	3.720^*	1.148	3.018	4.003^*
	SR×SL	0.372	1.986	3.116^*	1.704	5.388^**	5.487^**
茎	添加比（SR）	2.079	27.483^***	3.215	14.645^***	5.688^*	20.576^***
	添加量（SL）	3.859^*	3.171	1.106	3.115	0.484	3.718^*
	SR×SL	0.719	2.128	4.177^*	1.876	4.454^*	5.984^**
叶片	添加比（SR）	40.322^***	40.322^***	6.078^*	27.716^***	5.361^*	24.521^***
	添加量（SL）	0.360	4.076^*	4.112^*	3.949^*	4.087^*	2.478
	SR×SL	4.267^*	1.579	2.170	1.673	1.708	3.296^*
果实	添加比（SR）	0.964	0.612	10.709^**	15.620^***	11.660^**	28.414^***
	添加量（SL）	4.849	34.725^***	1.899	15.940^***	2.958	4.559^*
	SR×SL	0.280	4.145^*	6.031^**	4.462^*	5.138^**	4.594^*

氮磷添加处理	有机碳	全氮（TN）	全磷（TP）	碳氮比（C/N）	碳磷比（C/P）	氮磷比（N/P）
添加比（SR）	0.522	51.818^***	4.472^*	25.262^***	2.402	35.066^***
添加量（SL）	1.105	3.225	4.634^*	1.673	2.716	1.078
SR×SL	1.786	3.191^*	0.551	2.839	0.276	1.853