Response and correlation of above- and below-ground functional traits of Leymus chinensis to nitrogen and phosphorus additions

doi:10.17521/cjpe.2015.0164

Abstract

Abstract: AimsLeymus chinensis is a constructive and dominant species in typical steppe of northern China. The structure and functions of L. chinensis grassland ecosystem has been degenerated seriously due to long-term overgrazing in recent decades. As an effective measure to restore the degraded grasslands, the effects of nutrient addition on plant growth and ecosystem structure and functioning have been paid more attention in manipulation experimental research. The effects of nutrient addition, especially P addition on the above- and below-ground functional traits of L. chinensis have rarely been studied; particularly the underpinning mechanisms remain unclear. Our objective is to examine the responses and adaptive mechanisms of L. chinensis to different levels of N and P additions. MethodsWe conducted a culture experiment in the greenhouse, with three levels of N (50, 100 and 250 mg N·kg^-1) and P (5, 10 and 25 mg P·kg^-1) addition treatments. The above- and below-ground biomass, leaf traits (e.g., specific leaf area, leaf N and P contents) and root traits (e.g., specific root length, root N and P contents) of L. chinensis were determined in this study.Important findings Our results showed that: 1) the aboveground biomass and total biomass of L. chinensis were mostly affected by N addition, while the belowground biomass was mainly affected by P addition. N addition greatly enhanced the aboveground biomass of L. chinensis, while P addition reduced the belowground biomass at the moderate and high N levels. The root-shoot ratio of L. chinensis was influenced by both N and P additions, and root-shoot ratio decreased with increasing N and P levels. N and P additions promoted more biomass and N and P allocations to aboveground and leaf biomass. 2) Leymus chinensis showed different responses and adaptive mechanisms to P addition at low and high N levels. At low N level, L. chinensis exhibited high photosynthetic rate and specific root length (SRL) to improve photosynthetic capacity and root N acquisition, which promoted aboveground biomass. High root P content was favorable for belowground biomass. At high N level, P addition did not significantly affect plant growth of L. chinensis, even reduced its belowground biomass. Leymus chinensis showed high specific leaf area (SLA) and SRL to improve light interception and N acquisition in order to maintain stable aboveground biomass. 3) P addition greatly impacted below-ground than above-ground functional traits. SLA exhibited a weakly positive correlation with SRL, indicating L. chinensis exhibited relatively independence of resource acquirement and utilization between leaf and root functional traits.

http://jtp.cnki.net/bilingual/detail/html/ZWSB201601004

Key words: Leymus chinensis, plant functional traits, above- and below-ground relationships, nitrogen and phosphorus allocation, nitrogen and phosphorus additions

ZHAN Shu-Xia, ZHENG Shu-Xia, WANG Yang, BAI Yong-Fei. Response and correlation of above- and below-ground functional traits of Leymus chinensis to nitrogen and phosphorus additions[J]. Chin J Plant Ecol, 2016, 40(1): 36-47.

Add to citation manager EndNote|Ris|BibTeX

URL: https://www.plant-ecology.com/EN/10.17521/cjpe.2015.0164

https://www.plant-ecology.com/EN/Y2016/V40/I1/36

Figures/Tables 7

Fig. 1 Effects of N and P additions on individual aboveground biomass (A), belowground biomass (B), total biomass (C), and root: shoot ratio (D) of Leymus chinensis (mean ± SE). Different letters indicate significant difference (p < 0.05) among different P treatments at the same N level. p values indicate significant levels, and ns indicates non-significant difference among P treatments. N0P0, control; N1, N2, N3 represent low, moderate and high N levels, and P1, P2, P3 represent low, moderate and high P levels.

Fig. 2 Effects of N and P additions on N (A) and P (B) allocation between leaf and root biomass of Leymus chinensis (mean ± SE). N allocation is calculated as the ratio of leaf N biomass to root N biomass, and P allocation is the ratio of leaf P biomass to root P biomass. Different letters indicate significant difference (p < 0.05) among different P treatments at the same N level. p values indicate significant levels, and ns indicates non-significant difference among P treatments. N0P0, control; N1, N2, N3 represent low, moderate and high N levels, and P1, P2, P3 represent low, moderate and high P levels.

Fig. 3 Effects of N and P additions on leaf (A) and root (B) N, P, and N:P ratios of Leymus chinensis (mean ± SE). Different letters indicate significant difference (p < 0.05) among different P treatments at the same N level. P values indicate significant levels, and ns indicates non-significant difference among P treatments. N0P0, control; N1, N2, N3 represent low, moderate and high N levels, and P1, P2, P3 represent low, moderate and high P levels.

Fig. 4 Effects of N and P additions on specific leaf area (A) and specific root length (B) of Leymus chinensis (mean ± SE). Different letters indicate significant difference (p < 0.05) among different P treatments at the same N level. p values indicate significant levels, and ns indicates non-significant difference among P treatments. N0P0, control; N1, N2, N3 represent low, moderate and high N levels, and P1, P2, P3 represent low, moderate and high P levels.

Fig. 5 Relationships between individual aboveground and belowground biomass (A), specific leaf area and specific root length (B), leaf N and root N contents (C), leaf P and root P contents (D) of Leymus chinensis.

References 30

1	Bai X, Cheng JH, Zheng SX, Zhan SX, Bai YF (2014). Ecophysiological responses of Leymus chinensis to nitrogen and phosphorus additions in a typical steppe.Chinese Journal of Plant Ecology, 38, 103-115. (in Chinese with English abstract)
	[白雪, 程军回, 郑淑霞, 詹书侠, 白永飞 (2014). 典型草原建群种羊草对氮磷添加的生理生态响应. 植物生态学报, 38, 103-115.]
2	Bai YF, Wu JG, Clark CM, Naeem S, Pan QM, Huang JH, Zhang LX, Han XG (2010). Tradeoffs and thresholds in the effects of nitrogen addition on biodiversity and ecosystem functioning: Evidence from Inner Mongolia grasslands.Global Change Biology, 16, 358-372.
3	Chen SP, Bai YF, Zhang LX, Han XG (2005). Comparing physiological responses of two dominant grass species to nitrogen addition in Xilin River Basin of China.Environmental and Experimental Botany, 53, 65-75.
4	Cornelissen JHC, Lavorel S, Garnier E, Díaz S, Buchmann N, Gurvich DE, Reich PB, ter Steege H, Morgan HD, van der Heijden MGA, Pausas JG, Poorter H (2003). A handbook of protocols for standardised and easy measurement of plant functional traits worldwide.Australian Journal of Botany, 51, 335-380.
5	Elser JJ, Fagan WF, Denno RF, Dobberfuhl DR, Folarin A, Huberty A, Interlandi S, Kilham SS, McCauley E, Schulz KL, Siemann EH, Sterner RW (2000). Nutritional constraints in terrestrial and freshwater food webs.Nature, 408, 578-580.
6	Hart MM, Reader RJ, Klironomos JN (2003). Plant coexistence mediated by arbuscular mycorrhizal fungi.Trends in Ecology & Evolution, 18, 418-423.
7	Huang JY, Xu P, Yu HL, Yuan ZY, Li LH (2012). Responses of biomass, nutrient allocation of Leymus chinensis along N, P and water gradients.Pratacultural Science, 29, 1589-1595. (in Chinese with English abstract)
	[黄菊莹, 徐鹏, 余海龙, 袁志友, 李凌浩 (2012). 羊草生物量和养分分配对养分和水分添加的响应. 草业科学, 29, 1589-1595.]
8	Koerselman W, Meuleman AFM (1996). The vegetation N:P ratio: A new tool to detect the nature of nutrient limitation.Journal of Applied Ecology, 33, 1441-1450.
9	Lan ZC, Bai YF (2012). Testing mechanisms of N-enrichment- induced species loss in a semiarid Inner Mongolia grassland: Critical thresholds and implications for long-term ecosystem responses.Philosophical Transactions of the Royal Society B: Biological Sciences, 367, 3125-3134.
10	Lapointe BE (1987). Phosphorus-and nitrogen-limited photosynthesis and growth of Gracilaria tikvahiae (Rhodophyceae) in the Florida Keys: An experimental field study.Marine Biology, 93, 561-568.
11	Lavorel S, Grigulis K (2012). How fundamental plant functional trait relationships scale-up to trade-offs and synergies in ecosystem services.Journal of Ecology, 100, 128-140.
12	Lei Y, Hao ZP, Chen BD (2013). Effects of indigenous AM fungi and neighboring plants on the growth and phosphorus nutrition of Leymus chinensis.Acta Ecologica Sinica, 33, 1071-1079. (in Chinese with English abstract)
	[雷垚, 郝志鹏, 陈保冬 (2013). 土著菌根真菌和混生植物对羊草生长和磷营养的影响. 生态学报, 33, 1071-1079.]
13	Li XL, Feng G (2001). Eco-Physiology of Arbuscular Mycorrhiza. Sino-Culture Press, Beijing. 178-199.
14	Li YH (1993). Grazing dynamics of the species diversity in Aneurolepidium chinense steppe and Stipa grandis steppe.Acta Botanica Sinica, 35, 877-884. (in Chinese with English abstract)
	[李永宏 (1993). 放牧影响下羊草草原和大针茅草原植物多样性的变化. 植物学报, 35, 877-884.]
15	Liu GF, Freschet GT, Pan X, Cornelissen JHC, Li Y, Dong M (2010). Coordinated variation in leaf and root traits across multiple spatial scales in Chinese semi-arid and arid ecosystems.New Phytologist, 188, 543-553.
16	Lü XT, Reed S, Yu Q, He NP, Wang ZW, Han XG (2013). Convergent responses of nitrogen and phosphorus resorption to nitrogen inputs in a semiarid grassland.Global Change Biology, 19, 2775-2784.
17	Marklein AR, Houlton BZ (2012). Nitrogen inputs accelerate phosphorus cycling rates across a wide variety of terrestrial ecosystems.New Phytologist, 193, 696-704.
18	Müller I, Schmid B, Weiner J (2000). The effect of nutrient availability on biomass allocation patterns in 27 species of herbaceous plants.Perspectives in Plant Ecology, Evolution and Systematics, 3, 115-127.
19	Pan QM, Bai YF, Han XG, Yang JC (2005). Effects of nitrogen additions on a Leymus chinensis population in typical steppe of Inner Mongolia.Acta Phytoecologica Sinica, 29, 311-317. (in Chinese with English abstract)
	[潘庆民, 白永飞, 韩兴国, 杨景成 (2005). 氮素对内蒙古典型草原羊草种群的影响. 植物生态学报, 29, 311-317.]
20	Pan QM, Bai YF, Wu JG, Han XG (2011). Hierarchical plant responses and diversity loss after nitrogen addition: Testing three functionally-based hypotheses in the Inner Mongolia grassland.PLoS ONE, 6, e20078. doi: 10.1371/jour- nal.pone.0020078.
21	Reich PB, Oleksyn J, Wright IJ (2009). Leaf phosphorus influences the photosynthesis-nitrogen relation: A cross-biome analysis of 314 species.Oecologia, 160, 207-212.
22	Terry N, Ulrich A (1973). Effects of phosphorus deficiency on the photosynthesis and respiration of leaves of sugar beet.Plant Physiology, 51, 43-47.
23	Wan HW, Yang Y, Bai SQ, Xu YH, Bai YF (2008). Variations in leaf functional traits of six species along a nitrogen addition gradient in Leymus chinensis steppe in Inner Mongolia. Journal of Plant Ecology (Chinese Version), 32, 611-621. (in Chinese with English abstract)
	[万宏伟, 杨阳, 白世勤, 徐云虎, 白永飞 (2008). 羊草草原群落6种植物叶片功能特性对氮素添加的响应. 植物生态学报, 32, 611-621.]
24	Wang RZ (1997). The niche breaths and niche overlaps of main plant populations in Leymus chinensis grassland for grazing.Acta Phytoecologica Sinica, 21, 304-311. (in Chinese with English abstract)
	[王仁忠 (1997). 放牧影响下羊草草地主要植物种群生态位宽度与生态位重叠的研究. 植物生态学报, 21, 304-311.]
25	Wang YH, He XY, Zhou GS (2002). Study on the responses of Leymus chinensis steppe to grazing in Songnen Plain.Acta Agrestia Sinica, 10, 45-49. (in Chinese with English abstract)
	[王玉辉, 何兴元, 周广胜 (2002). 放牧强度对羊草草原的影响. 草地学报, 10, 45-49.]
26	Wang YL, Xu ZZ, Zhou GS (2004). Changes in biomass allocation and gas exchange characteristics of Leymus chinensis in response to soil water stress.Acta Phytoecologica Sinica, 28, 803-809. (in Chinese with English abstract)
	[王云龙, 许振柱, 周广胜 (2004). 水分胁迫对羊草光合产物分配及其气体交换特征的影响. 植物生态学报, 28, 803-809.]
27	Xu JC, Xu XY (2013). A study on growth rate and the growth rate hypothesis in Leymus chinensis.Pratacultural Science, 30, 74-79. (in Chinese with English abstract)
	[徐劲草, 许新宜 (2013). 羊草生长率的研究和生长率假说的验证. 草业科学, 30, 74-79.]
28	Xu ZZ, Zhou GS (2006). Combined effects of water stress and high temperature on photosynthesis, nitrogen metabolism and lipid peroxidation of a perennial grass Leymus chinensis.Planta, 224, 1080-1090.
29	Zhang LX, Bai YF, Han XG (2003). Application of N:P stoichiometry to ecology studies.Acta Botanica Sinica, 45, 1009-1018.
30	Zhang LX, Bai YF, Han XG (2004). Differential responses of N:P stoichiometry of Leymus chinensis and Carex korshinskyi to N additions in a steppe ecosystem in Nei Mongol.Acta Botanica Sinica, 46, 259-270.

[1]	DAI Jing-Zhong, BAI Yu-Ting, WEI Zhi-Jun, ZHANG Chu, YAN Rui-Rui. Effects of root-cutting in the vegetative phase on plant functional traits of Leymus chinensis [J]. Chin J Plant Ecol, 2021, 45(12): 1292-1302.
[2]	LI Ying, GONG Ji-Rui, LIU Min, HOU Xiang-Yang, DING Yong, YANG Bo, ZHANG Zi-He, WANG Biao, ZHU Chen-Chen. Defense strategies of dominant plants under different grazing intensity in the typical temperate steppe of Nei Mongol, China [J]. Chin J Plant Ecol, 2020, 44(6): 642-653.
[3]	ZHANG Lu, HAO Bi-Tai, QI Li-Xue, LI Yan-Long, XU Hui-Min, YANG Li-Na, BAOYIN Taogetao. Dynamic responses of aboveground biomass and soil organic matter content to grassland restoration [J]. Chin J Plant Ecol, 2018, 42(3): 317-326.
[4]	Xue YANG, Jun-Fang SHEN, Nian-Xi ZHAO, Yu-Bao GAO. Phenotypic plasticity and genetic differentiation of quantitative traits in genotypes of Leymus chinensis [J]. Chin J Plant Ecol, 2017, 41(3): 359-368.
[5]	Zhan-Wei ZHAI, Ji-Rui GONG, Qin-Pu LUO, Yan PAN, Taogetao BAOYIN, Sha XU, Min LIU, Li-Li YANG. Effects of nitrogen addition on photosynthetic characteristics of Leymus chinensis in the temperate grassland of Nei Mongol, China [J]. Chin J Plant Ecol, 2017, 41(2): 196-208.
[6]	Yu CEN, Mei-Zhen LIU. Effects of dew on eco-physiological traits and leaf structures of Leymus chinensis and Agropyron cristatum grown under drought stress [J]. Chin J Plan Ecolo, 2017, 41(11): 1199-1207.
[7]	LI Xi-Liang,HOU Xiang-Yang,WU Xin-Hong,null null,JI Lei,CHEN Hai-Jun,LIU Zhi-Ying,DING Yong. Plastic responses of stem and leaf functional traits in Leymus chinensis to long-term grazing in a meadow steppe [J]. Chin J Plant Ecol, 2014, 38(5): 440-451.
[8]	BAI Xue, CHENG Jun-Hui, ZHENG Shu-Xia, ZHAN Shu-Xia, BAI Yong-Fei. Ecophysiological responses of Leymus chinensis to nitrogen and phosphorus additions in a typical steppe [J]. Chin J Plant Ecol, 2014, 38(2): 103-115.
[9]	ZHOU Yong, ZHENG Lu-Yu, ZHU Min-Jie, LI Xia, REN An-Zhi, GAO Yu-Bao. Effects of fungal endophyte infection on soil properties and microbial communities in the host grass habitat [J]. Chin J Plant Ecol, 2014, 38(1): 54-61.
[10]	MA Hong-Yuan, LÜ Bing-Sheng, YANG Hao-Yu, YAN Chao, LIANG Zheng-Wei. Responses of seed germination of Leymus chinensis to environmental factors in degraded grassland on Songnen Plain in China [J]. Chin J Plant Ecol, 2012, 36(8): 812-818.
[11]	HUANG Ju-Ying, YU Hai-Long, YUAN Zhi-You, LI Ling-Hao. Effects of nitrogen, phosphorus and water supply on litter decomposition quality of senescing leaves of Leymus chinensis [J]. Chin J Plant Ecol, 2011, 35(8): 808-815.
[12]	GENG Yun-Xia, LI Yi-Ling, ZHU Sha, ZHU Jing-Jing, JIANG Jun-Cheng, NIU Hong-Hao, JIE Dong-Mei. Morphological changes of phytoliths in Leymus chinensis under saline-alkali stress [J]. Chin J Plant Ecol, 2011, 35(11): 1148-1155.
[13]	HAO Yan-Bin, WANG Yan-Fen, CUI Xiao-Yong. Drought stress reduces the carbon accumulation of the Leymus chinensis steppe in Inner Mongolia, China [J]. Chin J Plant Ecol, 2010, 34(8): 898-906.
[14]	WANG Xin-Ting, WANG Wei, LIANG Cun-Zhu. CHANGES IN THE POPULATION SPATIAL DISTRIBUTION PATTERN OF LEYMUS CHINENSIS IN DEGRADED STEPPE COMMUNITY DURING RESTORATIVE SUCCESSION IN INNER MONGOLIA, CHINA [J]. Chin J Plant Ecol, 2009, 33(1): 63-70.
[15]	WANG Yong-Fen, MO Xing-Guo, HAO Yan-Bin, GUO Rui-Ping, HUANG Xiang-Zhong, WANG Yan-Fen. SIMULATING SEASONAL AND INTERANNUAL VARIATIONS OF ECOSYS- TEM EVAPOTRANSPIRATION AND ITS COMPONENTS IN INNER MONG- OLIA STEPPE WITH VIP MODEL [J]. Chin J Plant Ecol, 2008, 32(5): 1052-1060.