糙隐子草功能性状对氮添加和干旱的响应

doi:10.17521/cjpe.2015.0004

摘要/Abstract

摘要：

植物功能性状被广泛地用于研究植物对环境变化的响应。糙隐子草(Cleistogenes squarrosa)是内蒙古草原重要的C₄物种, 其功能性状是如何对水氮环境的变化做出响应的, 还不十分清楚。该文采用盆栽实验的方法, 进行氮添加(0, 10.5, 35.0和56.0 g·m^-2·a^-1)和降水(自然降水和70%平均月降水量)处理, 研究糙隐子草整株性状、叶形态性状和叶生理性状对氮添加和干旱的响应。结果表明, 氮添加显著影响了糙隐子草的整株性状, 氮、水处理及它们的交互作用显著影响了糙隐子草的叶形态性状和叶生理性状。各功能性状对氮添加的响应格局在自然降水和干旱处理下是不同的。根深、茎生物量和茎叶比在干旱条件下低和中氮添加处理中较高, 而在自然降水下无明显变化; 比叶面积在干旱条件下随氮添加量的增加而增加, 而在自然降水下无增加趋势; 自然降水下, 高氮添加显著刺激了光合速率和蒸腾速率, 增加了水分利用效率, 而在干旱条件下氮添加对它们没有显著影响; 叶片单位面积的氮含量在自然降水下随氮添加量的增加有增加趋势, 而在干旱条件下显著降低。在自然降水下, 氮添加主要影响糙隐子草的叶形态和生理性状, 而在干旱条件下, 氮添加主要影响糙隐子草的整株性状和形态性状。总之, 糙隐子草的功能性状对氮添加表现出明显的响应, 响应格局在不同的水分条件下不同, 反映了其对氮水环境变化的弹性适应。

关键词: C4植物, 叶氮含量, 形态性状, 内蒙古草原, 光合速率, 生理性状

Abstract: Aims

Plant functional traits have been widely used to study the responses of plant to environmental change. Cleistogenes squarrosa is an important C₄ species in Nei Mongol grassland. How its functional traits would respond to varied nitrogen and water conditions have rarely been studied. Our subject was to examine the responses of the whole-plant traits and leaf morphological and physiological traits to nitrogen addition and drought in this species.

Methods

We conducted a pot experiment with a gradient of N addition (0, 10.5, 35.0, and 56.0 g·m^-2·a^-1) and water treatments (natural precipitation vs. 70% of mean monthly precipitation) in 2013. The whole-plant traits (e.g., root depth and stem-leaf biomass ratio), leaf morphological (e.g., leaf area and specific leaf area) and physiological traits (e.g., photosynthetic rate, water use efficiency, and leaf N content) were investigated.

<i>Important findings </i>

N addition had a significant effect on the whole-plant traits in C. squarrosa. The effects of N addition, water treatments, and an interaction between N addition and water treatments on leaf morphological and physiological traits were highly significant in most cases. The patterns of functional traits in response to N addition differed between plants under natural precipitation and with reduced mean monthly precipitation. Root depth, stem biomass, and stem-leaf biomass ratio were increased in treatments with low and intermediate N additions under reduced precipitation, but not changed under natural precipitation. Specific leaf area increased along the N addition gradient under drought, but did not change under natural precipitation. High N addition stimulated photosynthetic rate and transpiration rate and increased water use efficiency under natural precipitation, but had no effect on under reduced precipitation. Leaf N content on area basis increased slightly with the increases in N addition under natural precipitation, but decreased significantly under reduced precipitation. N addition influenced mainly the leaf morphological and physiological traits under natural precipitation and the whole-plant traits and leaf morphological traits under reduced precipitation. In conclusion, our results indicate that the functional traits in C. squarrosa respond to N addition and the patterns of responses differ under different water conditions, reflecting the adaptation to changes in N and water availability.

Key words: C4 plant, leaf nitrogen content, morphological traits, Nei Mongol grassland, photosynthetic rate, physiological traits

杨浩, 罗亚晨. 糙隐子草功能性状对氮添加和干旱的响应. 植物生态学报, 2015, 39(1): 32-42. DOI: 10.17521/cjpe.2015.0004

YANG Hao,LUO Ya-Chen. Responses of the functional traits in Cleistogenes squarrosa to nitrogen addition and drought. Chinese Journal of Plant Ecology, 2015, 39(1): 32-42. DOI: 10.17521/cjpe.2015.0004

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2015.0004

https://www.plant-ecology.com/CN/Y2015/V39/I1/32

图/表 8

图1 样地布局图。

Fig. 1 Plot layout.

表1 氮添加、水处理和氮水交互作用对糙隐子草整株性状和叶形态性状的影响

Table 1 Results of two-way ANOVAs for the effects of N addition, water treatments and their interaction on whole-plant traits and leaf morphological traits in Cleistogenes squarrosa

	整株性状 Whole-plant traits										叶形态性状 Leaf morphological traits
	高度 Height (cm)		根系深度 Root depth (cm)		茎生物量 Stem biomass (mg·tiller^-1)		叶生物量 Leaf biomass (mg·tiller^-1)		茎叶比 Stem: leaf ratio (tiller^-1)		叶面积 Leaf area (cm²·leaf^-1)		叶干质量 Leaf dry mass (mg·leaf^-1)		比叶面积 Specific leaf area (cm²·g^-1·leaf^-1)
	F	p	F	p	F	p	F	p	F	p	F	p	F	p	F	p
氮添加 N addition	4.930	0.010^*	15.340	0.000^**	16.940	0.000^**	6.080	0.004^**	12.040	0.000^**	2.940	0.059	0.000	0.986	3.540	0.035^*
水处理 Water treatment	0.000	1.000	0.000	1.000	0.000	1.000	0.000	1.000	0.000	1.000	2.870	0.106	3.680	0.031^*	5.480	0.030^*
氮添加×水处理 N addition × Water treatment	0.000	1.000	0.000	1.000	0.000	1.000	0.000	1.000	0.000	1.000	5.950	0.005^**	8.030	0.001^**	5.310	0.008^**

图2 施肥和干旱对糙隐子草整株性状的影响(平均值±标准误差, n = 4)。不同的字母表示处理间有显著的差异(LSD检验, p < 0.05)。

Fig. 2 Effects of N addition and drought on whole-plant traits in Cleistogenes squarrosa (mean ± SE, n = 4). Difference letters indicate significant differences among treatments (LSD test, p < 0.05).

表2 氮添加、水处理和氮水交互作用对糙隐子草叶生理性状和叶氮含量的影响

Table 2 Results of two-way ANOVAs for the effects of N addition, water treatments and their interaction on leaf photosynthetic traits and leaf nitrogen content in Cleistogenes squarrosa

	叶生理形状 Leaf photosynthetic traits													叶片氮含量 Leaf nitrogen content
	光合速率 P_n (μmol CO₂·m^-2·s^-1)		蒸腾速率 T_r (mmol H₂O·m^-2·s^-1)		气孔导度 G_s (mol H₂O·m^-2·s^-1)		胞间CO₂浓度 C_i (μmol CO₂·mol^-1)		水分利用效率 WUE (μmol CO₂·mol^-1H₂O)		气孔限制 L_s			单位质量叶片氮含量 N_mass(%)		单位面积叶片氮含量 N_area(g·m^-2)
	F	p	F	p	F	p	F	p	F	p	F	p		F	p	F	p
	氮添加 N addition	3.91	0.024^*	1.86	0.169	1.39	0.274	11.28	0.000^**	2.44	0.095	11.49	0.000^**		6.71	0.003^**	1.34	0.290
水处理 Water treatment	9.19	0.007^**	12.89	0.002^**	10.85	0.004^**	6.29	0.021^*	0.44	0.513	6.50	0.019^*		7.55	0.012^*	15.05	0.001**
氮添加×水处理 N addition × Water treatment	7.82	0.001^**	2.70	0.073	4.14	0.020^*	10.03	0.000^**	3.71	0.029^*	9.91	0.000^**		0.45	0.720	5.16	0.008**

图3 施肥和干旱对糙隐子草叶片形态性状的影响(平均值±标准误差, n = 4)。不同的字母表示处理间有显著的差异(LSD检验, p < 0.05)。

Fig. 3 Effects of N addition and drought on leaf morphological traits in Cleistogenes squarrosa (mean ± SE, n = 4). Difference letters indicate significant differences among treatments (LSD test, p < 0.05).

图4 施肥和干旱对糙隐子草叶片光合性状的影响(平均值±标准误差, n = 4)。Ci, 胞间CO2浓度; Gs, 气孔导度; Ls, 气孔限制值; Pn, 光合速率; Tr, 蒸腾速率; WUE, 水分利用效率。不同的字母表示处理间有显著的差异(LSD test, p < 0.05)。

Fig. 4 Effects of N addition and drought on leaf photosynthetic traits in Cleistogenes squarrosa (mean ± SE, n = 4). Ci, intercellular CO2 concentration; Gs, stomatal conductance; Ls, stomatal limitation; Pn, net photosynthetic rate; Tr, transpiration rate; WUE, water use efficiency. Difference letters indicate significant differences among treatments (LSD test, p < 0.05).

图5 施肥和干旱对糙隐子草叶片单位质量氮含量和单位面积氮含量的影响(平均值±标准误差, n = 4)。不同的字母表示处理间有显著的差异(LSD test, p < 0.05)。

Fig. 5 Effects of N addition and drought on leaf nitrogen content based on leaf mass (Nmass) and leaf nitrogen content based on leaf area (Narea) in Cleistogenes squarrosa (mean ± SE, n = 4). Difference letters indicate significant differences among treatments (LSD test, p < 0.05).

图6 性状间关系的聚类分析图。Ci, 胞间CO2浓度; Gs, 气孔导度; Ls, 气孔限制值; Narea, 单位面积叶片氮含量; Nmass, 单位质量叶片氮含量; Pn, 光合速率; Tr, 蒸腾速率; WUE, 水分利用效率。

Fig. 6 The cluster dendrogram of plant traits. Ci, intercellular CO2 concentration; Gs, stomatal conductance; Ls, stomatal limitation; Narea, leaf nitrogen content based on leaf area; Nmass, leaf nitrogen content based on leaf mass; Pn, net photosynthetic rate; Tr, transpiration rate; WUE, water use efficiency.

参考文献 29

1	Adler PB, Milchunas DG, Lauenroth WK, Sala OE, Burke IC (2004). Functional traits of graminoids in semi-arid steppes: A test of grazing histories. Journal of Applied Ecology, 41, 653-663.
2	Adler PB, Milchunas DG, Sala OE, Burke IC, Lauenroth WK (2005). Plant traits and ecosystem grazing effects: Comparison of U.S. Sagebrush steppe and Patagonian steppe. Ecological Applications, 15, 774-792.
3	Auyeung DSN, Suseela V, Dukes JS (2013). Warming and drought reduce temperature sensitivity of nitrogen transformations. Global Change Biology, 19, 662-676.
4	Bai YF, Han XG, Wu JG, Chen ZZ, Li LH (2004). Ecosystem stability and compensatory effects in the Inner Mongolia grassland. Nature, 431, 181-184.
5	Bai YF, Wu JG, Xing Q, Pan QM, Huang JH, Yang DL, Han XG (2008). Primary production and rain use efficiency across a precipitation gradient on the Mongolia Plateau. Ecology, 89, 2140-2153.
6	Chen SP, Bai YF, Han XG (2002). Variation of water-use efficiency of Leymus chinensis and Cleistogenes squarrosa in different plant communities in Xilin River Basin, Nei Mongol. Acta Botanica Sinica, 44, 1484-1490.
7	Chen SP, Bai YF, Lin GH, Liang Y, Han XG (2005). Effects of grazing on photosynthetic characteristics of major steppe species in the Xilin River Basin, Inner Mongolia, China. Photosynthetica, 43, 559-565.
8	Cui XY, Chen ZZ, Du ZC (2001). Study on light- and water-use characteristics of main plants in semiarid steppe. Acta Prataculturae Sinica, 10(2), 14-21.
	(in Chinese with English abstract) [崔骁勇, 陈佐忠, 杜占池 (2001). 半干旱草原主要植物光能和水分利用特征的研究. 草业学报, 10(2), 14-21.]
9	Díaz S, Lavorel S, McIntyre S, Falczuk V, Casanoves F, Milchunas DG, Skarpe C, Rusch G, Sternberg M, Noy-Meir I, Landsberg J, Zhang W, Clark H, Campbell BD (2007). Plant trait responses to grazing — A global synthesis. Global Change Biology, 13, 313-341.
10	Huang JY, Zhu XG, Yuan ZY, Song SH, Li X, Li LH (2008). Changes in nitrogen resorption traits of six temperate grassland species along a multi-level N addition gradient. Plant and Soil, 306, 149-158.
11	Inner Mongolia-Ningxia Complex Expert Team of the Chinese Academy of Sciences (1985). Vegetation of Inner Mongolia. Science Press, Beijing.
	(in Chinese) [中国科学院内蒙古宁夏综合考察队 (1985). 内蒙古植被. 科学出版社, 北京.]
12	Lavorel S, Grigulis K, Lamarque P, Colace MP, Garden D, Girel J, Pellet G, Douzet R (2011). Using plant functional traits to understand the landscape distribution of multiple ecosystem services. Journal of Ecology, 99, 135-147.
13	Lü CQ, Tian HQ (2007). Spatial and temporal patterns of nitrogen deposition in China: Synthesis of observational data. Journal of Geophysical Research: Atmospheres, 112, D22S05, doi: 10.1029/2006JD007990.
14	Milchunas DG, Lauenroth WK (1993). Quantitative effects of grazing on vegetation and soils over a global range of environments. Ecological Monographs, 63, 327-366.
15	Milla R, Reich PB (2007). The scaling of leaf area and mass: the cost of light interception increases with leaf size. Proceedings of the Royal Society B: Biological Sciences, 274, 2109-2115.
16	Paul KI, Polglase PJ, O’Connell AM, Carlyle JC, Smethurst PJ, Khanna PK (2003). Defining the relation between soil water content and net nitrogen mineralization. European Journal of Soil Science, 54, 39-48.
17	Poorter L, Rozendaal DA (2008). Leaf size and leaf display of thirty-eight tropical tree species. Oecologia, 158, 35-46.
18	Qin J, Bao YJ, Li ZH, Hu ZC, Zhou LN, Sun Z (2014). Gradient response of Cleistogenes squarrosa root system to nitrogen addition. Journal of Dalian Nationalities University, 16, 24-28.
	(in Chinese with English abstract) [秦洁, 鲍雅静, 李政海, 胡志超, 周丽娜, 孙振 (2014). 糙隐子草根系特征对氮素添加梯度的响应. 大连民族学院学报, 16, 24-28.]
19	Reich PB, Walters MB, Ellsworth DS (1997). From tropics to tundra: global convergence in plant functioning. Proceedings of the National Academy of Sciences of the United States of America, 94, 13730-13734.
20	Reich PB, Wright IJ, Lusk CH (2007). Predicting leaf physiology from simple plant and climate attributes: A global GLOPNET analysis. Ecological Applications, 17, 1982-1988.
21	Shipley B (2002). Trade-offs between net assimilation rate and specific leaf area in determining relative growth rate: Relationship with daily irradiance. Functional Ecology, 16, 682-689.
22	Shipley B (2006). Net assimilation rate, specific leaf area and leaf mass ratio: Which is most closely correlated with relative growth rate? A meta-analysis. Functional Ecology, 20, 565-574.
23	Violle C, Navas ML, Vile D, Kazakou E, Fortunel C, Hummel I, Garnier E (2007). Let the concept of trait be functional!Oikos, 116, 882-892.
24	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.]
25	Wang SP, Wang YF (2001). Study on over-compensation growth of Cleistogenes squarrosa population in Inner Mongolia steppe. Acta Batanica Sinica, 43, 413-418.
	(in Chinese with English abstract) [汪诗平, 王艳芬 (2001). 不同放牧率下糙隐子草种群补偿性生长的研究植物学报, 43, 413-418.]
26	Wang SP, Wang YF, Chen ZZ (2003). Effect of climate change and grazing on populations of Cleistogenes squarrosa in Inner Mongolia steppe. Acta Phytoecologica Sinica, 23, 337-343.
	(in Chinese with English abstract). [汪诗平, 王艳芬, 陈佐忠 (2003). 气候变化和放牧活动对糙隐子草种群的影响植物生态学报, 23, 337-343.]
27	Wang XT, Wang W, Liang CZ, Bao JJ (2014). Cleistogenes squarrosa population at different restorative succession stages in Inner Mongolia of China: A point pattern analysis. Chinese Journal of Applied Ecology, 24, 1793-1800.
	(in Chinese with English abstract) [王鑫厅, 王炜, 梁存柱, 包俊江 (2014). 不同恢复演替阶段糙隐子草种群的点格局分析. 应用生态学报, 24, 1793-1800.]
28	Westoby M, Falster DS, Moles AT, Vesk PA, Wright IJ (2002). Plant ecological strategies: Some leading dimensions of variation between species. Annual Review of Ecology and Systematics, 33, 125-159.
29	Zheng SX, Lan ZC, Li WH, Shao RX, Shan YM, Wan HW, Taube F, Bai YF (2011). Differential responses of plant functional trait to grazing between two contrasting dominant C3 and C4 species in a typical steppe of Inner Mongolia, China. Plant and Soil, 340, 141-155.

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