Effects of nitrogen and water addition on leaf nitrogen and phosphorus stoichiometry of the dominant species in an Artemisia ordosica community

doi:10.17521/cjpe.2022.0458

Abstract

Abstract:

Aims Water and nitrogen (N) availability are two of crucial factors affecting the growth of desert plants, and are sensitive to global climate change. With increasing N deposition and precipitation, the understanding of the responses of leaf N and phosphorus (P) stoichiometry of dominant species in sandy semi-shrub community could provide insights into the adaptive strategies of sand-fixing plants in the Mau Us Sandy Land.

Methods An experiment of N and water addition was conducted in an Artemisia ordosica community in the southern edge of the Mau Us Sandy Land. Soil available N and P concentrations, soil available N:P, leaf N and P concentrations, leaf N:P, and the corresponding homeostasis index of dominant species A. ordosica and Leymus secalinus were measured.

Important findings (1) Soil N and P availability and species identity jointly influenced the response of leaf N concentration, leaf P concentration, and leaf N:P in A. ordosica and L. secalinus to N and water addition, and the leaf N concentration of A. ordosica and L. secalinus showed different responses to treatments. (2) Both A. ordosica and L. secalinus showed higher stoichiometric homoeostasis indices for leaf P than for leaf N, which was closely related to the fact that the growth of A. ordosica and L. secalinuswere both N-limited. (3) The stoichiometric homoeostasis indices of leaf N concentration, leaf P concentration, and N:P in A. ordosica were relatively higher than those of L. secalinus, and A. ordosica adopted more conservative nutrient utilization strategies, while L. secalinus adopted more flexible nutrient utilization strategies. Therefore, A. ordosica was more competitive than L. secalinus in growth in arid and infertile environments. Under the scenarios of increasing N deposition and precipitation, the species composition of the A. ordosica community may change due to the contrasting competitiveness and nutrient utilization strategies between A. ordosica and L. secalinus.

Key words: Mau Us Sandy Land, nitrogen deposition, precipitation alteration, ecological stoichiometry, nutrient limitation, homoeostasis

ZHANG Wen-Jin, SHE Wei-Wei, QIN Shu-Gao, QIAO Yan-Gui, ZHANG Yu-Qing. Effects of nitrogen and water addition on leaf nitrogen and phosphorus stoichiometry of the dominant species in an Artemisia ordosica community[J]. Chin J Plant Ecol, 2024, 48(5): 590-600.

Add to citation manager EndNote|Ris|BibTeX

URL: https://www.plant-ecology.com/EN/10.17521/cjpe.2022.0458

https://www.plant-ecology.com/EN/Y2024/V48/I5/590

Figures/Tables 7

Fig. 1 Changes of soil moisture under nitrogen (N) and water addition treatments (mean ± SE). **, p < 0.01; ***, p < 0.001; ns, p > 0.05. Different lowercase letters indicate significant differences among different N addition treatments of the same water addition treatment (p < 0.05), and different uppercase letters indicate significant differences among different water addition treatments (p < 0.05). N, effects of N addition; W, effects of water addition; N × W, interactive effects of N and water addition. N0, N05, N10, N20, N30, and N60 represent nitrogen addition amounts of 0, 0.5, 1.0, 2.0, 3.0, and 6.0 g·m-2·a-1, respectively. W0, ambient precipitation; W20, increased precipitation by 20%; W40, increased precipitation by 40%.

Fig. 2 Changes of soil available nitrogen (N) concentrations, available phosphorus (P) concentrations, and available N:P under N and water addition treatments (mean ± SE). *, p < 0.05; ***, p < 0.001; ns, p > 0.05. Different lowercase letters indicate significant differences among different N addition treatments of the same water addition treatment (p < 0.05), and different uppercase letters indicate significant differences among different water addition treatments (p < 0.05). N, effects of N addition; W, effects of water addition; N × W, interactive effects of N and water addition. N0, N05, N10, N20, N30, and N60 represent nitrogen addition amounts of 0, 0.5, 1.0, 2.0, 3.0, and 6.0 g·m-2·a-1, respectively. W0, ambient precipitation; W20, increased precipitation by 20%; W40, increased precipitation by 40%.

Fig. 3 Changes of leaf nitrogen (N) and phosphorus (P) stoichiometry of Artemisia ordosica and Leymus secalinus under N and water addition treatments (mean ± SE). *, p < 0.05; **, p < 0.01; ***, p < 0.001; ns, p > 0.05. Different lowercase letters indicate significant differences among different N addition treatments of the same water addition treatment (p < 0.05), and different uppercase letters indicate significant differences among different water addition treatments (p < 0.05). N, effects of N addition; W, effects of water addition; N × W, interactive effects of N and water addition. N0, N05, N10, N20, N30, and N60 represent nitrogen addition amounts of 0, 0.5, 1.0, 2.0, 3.0, and 6.0 g·m-2·a-1, respectively. W0, ambient precipitation; W20, increased precipitation by 20%; W40, increased precipitation by 40%.

Fig. 4 Comparisons of leaf nitrogen (N) and phosphorus (P) stoichiometry between two dominant species Artemisia ordosica and Leymus secalinus (mean ± SE). Different lowercase letters indicate significant differences between species (p < 0.05). CV, coefficient of variation.

Fig. 5 Relationships of leaf nitrogen (N) concentration, leaf phosphorus (P) concentration, and leaf N:P with soil available N concentration, soil available P concentration, and soil available N:P. HN, HP, and HN:P represent the homeostasis indices of leaf N concentration, leaf P concentration, and leaf N:P, respectively.

Fig. 6 Changes in the coverage of dominant species Artemisia ordosica and Leymus secalinus under nitrogen (N) and water addition treatments (mean ± SE). **, p < 0.01; ***, p < 0.001; ns, p > 0.05. Different lowercase letters indicate significant differences among different N addition treatments of the same water addition treatment (p < 0.05), and different uppercase letters indicate significant differences among different water addition treatments (p < 0.05). N, effects of N addition; W, effects of water addition; N × W, interactive effects of N and water addition. N0, N05, N10, N20, N30, and N60 represent nitrogen addition amounts of 0, 0.5, 1.0, 2.0, 3.0, and 6.0 g·m-2·a-1, respectively. W0, ambient precipitation; W20, increased precipitation by 20%; W40, increased precipitation by 40%.

Table 1 Correlation coefficients between soil available nitrogen (N), soil available phosphorus (P) concentrations and leaf N, leaf P concentrations of Artemisia ordosica and Leymus secalinus

	叶片N含量 Leaf N concentration		叶片P含量 Leaf P concentration
	黑沙蒿 A. ordosica	赖草 L. secalinus	黑沙蒿 A. ordosica	赖草 L. secalinus
土壤速效N含量 Soil available N concentration	0.507^***	0.419^**	-	-
土壤速效P含量 Soil available P concentration	-	-	-0.074	0.030

References 50

[1]	Chen CD (2009). Towards Macroecology: an Anthology of Chen Changdu’s Papers. Science Press, Beijing.
	[陈昌笃 (2009). 走向宏观生态学: 陈昌笃论文选集. 科学出版社, 北京.]
[2]	Chen Y, Li YQ, Wang XY, Niu YY (2021). Advances in ecological stoichiometry in typically and ecologically vulnerable regions of China. Acta Ecologica Sinica, 41, 4213-4225.
	[陈云, 李玉强, 王旭洋, 牛亚毅 (2021). 中国典型生态脆弱区生态化学计量学研究进展. 生态学报, 41, 4213-4225.]
[3]	Cross AF, Schlesinger WH (2001). Biological and geochemical controls on phosphorus fractions in semiarid soils. Biogeochemistry, 52, 155-172.
[4]	Deng Q, Hui D, Dennis S, Reddy KC (2017). Responses of terrestrial ecosystem phosphorus cycling to nitrogen addition: a meta-analysis. Global Ecology and Biogeography, 26, 713-728.
[5]	Dijkstra FA, Pendall E, Morgan JA, Blumenthal DM, Carrillo Y, LeCain DR, Follett RF, Williams DG (2012). Climate change alters stoichiometry of phosphorus and nitrogen in a semiarid grassland. New Phytologist, 196, 807-815. DOI PMID
[6]	Elser JJ, Bracken MES, Cleland EE, Gruner DS, Harpole WS, Hillebrand H, Ngai JT, Seabloom EW, Shurin JB, Smith JE (2007). Global analysis of nitrogen and phosphorus limitation of primary producers in freshwater, marine and terrestrial ecosystems. Ecology Letters, 10, 1135-1142. DOI PMID
[7]	Elser JJ, Fagan WF, Kerkhoff AJ, Swenson NG, Enquist BJ (2010). Biological stoichiometry of plant production: metabolism, scaling and ecological response to global change. New Phytologist, 186, 593-608. DOI PMID
[8]	Galloway JN, Dentener FJ, Capone DG, Boyer EW, Howarth RW, Seitzinger SP, Asner GP, Cleveland CC, Green PA, Holland EA, Karl DM, Michaels AF, Porter JH, Townsend AR, Vöosmarty CJ (2004). Nitrogen cycles: past, present, and future. Biogeochemistry, 70, 153-226.
[9]	Garnier E (1998). Interspecific variation in plasticity of grasses in response to nitrogen supply//Cheplick GP. Population Biology of Grasses. Cambridge University Press, Cambridge, UK. 155-182.
[10]	Huang G, Liu Y, Huang RH (2011). The interannual variability of summer rainfall in the arid and semiarid regions of Northern China and its association with the Northern Hemisphere circumglobal teleconnection. Advances in Atmospheric Sciences, 28, 257-268.
[11]	Huang JY, Yu HL, Liu JL, Ma F, Han L (2018). Effects of precipitation levels on the C:N:P stoichiometry in plants, microbes, and soils in a desert steppe in China. Acta Ecologica Sinica, 38, 5362-5373.
	[黄菊莹, 余海龙, 刘吉利, 马飞, 韩磊 (2018). 控雨对荒漠草原植物、微生物和土壤C、N、P化学计量特征的影响. 生态学报, 38, 5362-5373.]
[12]	Jiang LL, Zeng CS, Shao JJ, Zhou XH (2017). Plant nutrient dynamics and stoichiometric homeostasis of invasive species Spartina alterniflora and native Cyperus malaccensis var. brevifolius in the Minjiang River estuarine wetlands. Chinese Journal of Plant Ecology, 41, 450-460.
	[蒋利玲, 曾从盛, 邵钧炯, 周旭辉 (2017). 闽江河口入侵种互花米草和本地种短叶茳芏的养分动态及植物化学计量内稳性特征. 植物生态学报, 41, 450-460.] DOI
[13]	Lai ZR, Zhang YQ, Liu JB, Wu B, Qin SG, Fa KY (2016). Fine-root distribution, production, decomposition, and effect on soil organic carbon of three revegetation shrub species in northwest China. Forest Ecology and Management, 359, 381-388.
[14]	Li YL, Mao W, Zhao XY, Zhang TH (2010). Leaf nitrogen and phosphorus stoichiometry in typical desert and desertified regions, North China. Environmental Science, 31, 1716-1725.
	[李玉霖, 毛伟, 赵学勇, 张铜会 (2010). 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. 环境科学, 31, 1716-1725.]
[15]	Liu JB, Fa KY, Zhang YQ, Wu B, Qin SG, Jia X (2015). Abiotic CO₂ uptake from the atmosphere by semiarid desert soil and its partitioning into soil phases. Geophysical Research Letters, 42, 5779-5785.
[16]	Liu L, Bai YX, Qiao YG, Miao C, She WW, Qin SG, Zhang YQ (2023). Water-use characteristics of two dominant plant species in different community types in the Mu Us Desert. Catena, 221, 106803. DOI: 10.1016/j.catena.2022. 106803.
[17]	Lü XT, Dijkstra FA, Kong DL, Wang ZW, Han XG (2014). Plant nitrogen uptake drives responses of productivity to nitrogen and water addition in a grassland. Scientific Reports, 4, 4817. DOI: 10.1038/srep04817.
[18]	Lü XT, Han XG (2010). Nutrient resorption responses to water and nitrogen amendment in semi-arid grassland of Inner Mongolia, China. Plant and Soil, 327, 481-491.
[19]	Lü XT, Hu YY, Zhang HY, Wei HW, Hou SL, Yang GJ, Liu ZY, Wang XB (2018). Intraspecific variation drives community-level stoichiometric responses to nitrogen and water enrichment in a temperate steppe. Plant and Soil, 423, 307-315.
[20]	Lü X, Kong D, Pan Q, Simmons ME, Han X (2012). Nitrogen and water availability interact to affect leaf stoichiometry in a semi-arid grassland. Oecologia, 168, 301-310.
[21]	Lü XT, Reed S, Hou SL, Hu YY, Wei HW, Lü FM, Cui Q, Han XG (2017). Temporal variability of foliar nutrients: responses to nitrogen deposition and prescribed fire in a temperate steppe. Biogeochemistry, 133, 295-305.
[22]	Luo X, Wang YT, Zhang J, Li JP (2022). Responses of dominant species and rhizosphere soil stoichiometry to rainfall in typical steppe of the Loess Plateau. Acta Ecologica Sinica, 42, 1002-1014.
	[罗叙, 王誉陶, 张娟, 李建平 (2022). 黄土高原典型草原优势种植物及其根际土壤化学计量对降雨变化的响应. 生态学报, 42, 1002-1014.]
[23]	Ma Y, Xu ZH, Zeng QH, Meng JL, Hu YH, Su JQ (2021). Impact of nitrogen addition on stoichiometric characteristics of herbaceous species in desert steppe. Acta Prataculturae Sinica, 30, 64-72. DOI
	[马英, 许志豪, 曾巧红, 孟建龙, 胡亚虎, 苏洁琼 (2021). 氮素添加对荒漠化草原草本植物养分化学计量特征的影响. 草业学报, 30, 64-72.] DOI
[24]	Mariotte P, Canarini A, Dijkstra FA (2017). Stoichiometric N: P flexibility and mycorrhizal symbiosis favour plant resistance against drought. Journal of Ecology, 105, 958-967.
[25]	Marklein AR, Houlton BZ (2012). Nitrogen inputs accelerate phosphorus cycling rates across a wide variety of terrestrial ecosystems. New Phytologist, 193, 696-704. DOI PMID
[26]	Ning ZY, Li YL, Yang HL, Zhang ZQ (2019). Nitrogen and phosphorus stoichiometric homoeostasis in leaves of dominant sand-fixing shrubs in Horqin Sandy Land, China. Chinese Journal of Plant Ecology, 43, 46-54.
	[宁志英, 李玉霖, 杨红玲, 张子谦 (2019). 科尔沁沙地优势固沙灌木叶片氮磷化学计量内稳性. 植物生态学报, 43, 46-54.] DOI
[27]	Perring MP, Hedin LO, Levin SA, McGroddy M, de Mazancourt C (2008). Increased plant growth from nitrogen addition should conserve phosphorus in terrestrial ecosystems. Proceedings of the National Academy of Sciences of the United States of America, 105, 1971-1976. DOI PMID
[28]	Persson J, Fink P, Goto A, Hood JM, Jonas J, Kato S (2010). To be or not to be what you eat: regulation of stoichiometric homeostasis among autotrophs and heterotrophs. Oikos, 119, 741-751.
[29]	Ren HY, Kang J, Yuan ZY, Xu ZW, Han GD (2018). Responses of nutrient resorption to warming and nitrogen fertilization in contrasting wet and dry years in a desert grassland. Plant and Soil, 432, 65-73.
[30]	Ren HY, Xu ZW, Huang JH, Lü XT, Zeng DH, Yuan ZY, Han XG, Fang YT (2015). Increased precipitation induces a positive plant-soil feedback in a semi-arid grassland. Plant and Soil, 389, 211-223.
[31]	Sardans J, Grau O, Chen HYH, Janssens IA, Ciais P, Piao S, Peñuelas J (2017). Changes in nutrient concentrations of leaves and roots in response to global change factors. Global Change Biology, 23, 3849-3856. DOI PMID
[32]	She WW, Bai YX, Zhang YQ, Qin SG, Feng W, Lai ZR, Qiao YG, Liu L, Zhang WJ, Miao C (2021). Nitrogen-enhanced herbaceous competition threatens woody species persistence in a desert ecosystem. Plant and Soil, 460, 333-345.
[33]	She WW, Bai YX, Zhang YQ, Qin SG, Jia X, Feng W, Lai ZR, Fu J, Qiao YG (2020). Nitrogen enrichment suppresses revegetated shrub growth under increased precipitation via herb-induced topsoil water limitation in a desert ecosystem in Northern China. Plant and Soil, 446, 97-110.
[34]	Song YF, Lu YJ, Liu TJ, Liu HW, Yan ZY, Wang HQ (2020). Soil-plant-microbial C, N, P and their stoichiometric characteristics in different rainfall zones of desert steppe. Acta Ecologica Sinica, 40, 4011-4023.
	[宋一凡, 卢亚静, 刘铁军, 刘慧文, 闫泽宇, 王慧琪 (2020). 荒漠草原不同雨量带土壤-植物-微生物C、N、P及其化学计量特征. 生态学报, 40, 4011-4023.]
[35]	Sterner RW, Elser JJ (2002). Ecological Stoichiometry—The Biology of Elements from Molecules to the Biosphere. Princeton University Press, Princeton.
[36]	Tang ZY, Xu WT, Zhou GY, Bai YF, Li JX, Tang XL, Chen DM, Liu Q, Ma WH, Xiong GM, He HL, He NP, Guo YP, Guo Q, Zhu JL, et al. (2018). Patterns of plant carbon, nitrogen, and phosphorus concentration in relation to productivity in China’s terrestrial ecosystems. Proceedings of the National Academy of Sciences of the United States of America, 115, 4033-4038.
[37]	Tian D, Yan ZB, Fang JY (2021). Review on characteristics and main hypotheses of plant ecological stoichiometry. Chinese Journal of Plant Ecology, 45, 682-713. DOI
	[田地, 严正兵, 方精云 (2021). 植物生态化学计量特征及其主要假说. 植物生态学报, 45, 682-713.] DOI
[38]	Tian D, Reich PB, Chen HYH, Xiang Y, Luo Y, Shen Y, Meng C, Han W, Niu S (2019). Global changes alter plant multi-element stoichiometric coupling. New Phytologist, 221, 807-817. DOI PMID
[39]	Trenberth KE (2011). Changes in precipitation with climate change. Climate Research, 47, 123-138.
[40]	Wang B, Huang G, Ma J, Li Y (2016). Responses of nutrients resorption of five desert ephemeral plants to water and nitrogen additions. Journal of Desert Research, 36, 415-422. DOI
	[王斌, 黄刚, 马健, 李彦 (2016). 5种荒漠短命植物养分再吸收对水氮添加的响应. 中国沙漠, 36, 415-422.] DOI
[41]	Wang R, Creamer CA, Wang X, He P, Xu Z, Jiang Y (2016). The effects of a 9-year nitrogen and water addition on soil aggregate phosphorus and sulfur availability in a semi-arid grassland. Ecological Indicators, 61, 806-814.
[42]	Xu W, Luo XS, Pan YP, Zhang L, Tang AH, Shen JL, Zhang Y, Li KH, Wu QH, Yang DW, Zhang YY, Xue J, Li WQ, Li QQ, Tang L, et al. (2015). Quantifying atmospheric nitrogen deposition through a nationwide monitoring network across China. Atmospheric Chemistry and Physics, 15, 12345-12360.
[43]	Yan ZB, Tian D, Han WX, Tang ZY, Fang JY (2017). An assessment on the uncertainty of the nitrogen to phosphorus ratio as a threshold for nutrient limitation in plants. Annals of Botany, 120, 937-942. DOI PMID
[44]	Yu Q, Chen Q, Elser JJ, He N, Wu H, Zhang G, Wu J, Bai Y, Han X (2010). Linking stoichiometric homoeostasis with ecosystem structure, functioning and stability. Ecology Letters, 13, 1390-1399. DOI PMID
[45]	Yu Q, Elser JJ, He N, Wu H, Chen Q, Zhang G, Han X (2011). Stoichiometric homeostasis of vascular plants in the Inner Mongolia grassland. Oecologia, 166, 1-10. DOI PMID
[46]	Yu Q, Wilcox K, La Pierre K, Knapp AK, Han X, Smith MD (2015). Stoichiometric homeostasis predicts plant species dominance, temporal stability, and responses to global change. Ecology, 96, 2328-2335. PMID
[47]	Zhang RY, Shi XM, Li WJ, Guo R, Wang G (2015). Response of species homeostasis and biomass on a sub-alpine grassland. Pratacultural Science, 32, 1539-1547.
	[张仁懿, 史小明, 李文金, 郭睿, 王刚 (2015). 亚高寒草甸物种内稳性与生物量变化模式. 草业科学, 32, 1539-1547.]
[48]	Zhang WJ, Zhang YQ, She WW, Qin SG, Feng W (2016). Effects of nitrogen addition on foliar ecological stoichiometric characteristics of Artemisia ordosica community. Research of Environmental Sciences, 29, 52-58.
	[张文瑾, 张宇清, 佘维维, 秦树高, 冯薇 (2016). 氮添加对油蒿群落植物叶片生态化学计量特征的影响. 环境科学研究, 29, 52-58.]
[49]	Zhang ZS, Yang GS, Lü XY, Hu R, Huang L (2022). Research progresses in ecological stoichiometry of C, N and P in desert ecosystems. Journal of Desert Research, 42, 48-56.
	[张志山, 杨贵森, 吕星宇, 虎瑞, 黄磊 (2022). 荒漠生态系统C、N、P生态化学计量研究进展. 中国沙漠, 42, 48-56.] DOI
[50]	Zhu WW, Xu YX, Wang P, Yu HL, Huang JY (2020). Effects of precipitation and N addition on plant and microbial C:N:P ecological stoichiometry in a desert steppe of northwestern China. Acta Botanica Boreali-Occidentalia Sinica, 40, 676-687.
	[朱湾湾, 许艺馨, 王攀, 余海龙, 黄菊莹 (2020). 降水量及N添加对荒漠草原植物和土壤微生物C:N:P生态化学计量特征的影响. 西北植物学报, 40, 676-687.]