长期氮添加对典型草原植物多样性与初级生产力的影响及途径

doi:10.17521/cjpe.2019.0260

植物生态学报 ›› 2020, Vol. 44 ›› Issue (1): 22-32.DOI: 10.17521/cjpe.2019.0260

长期氮添加对典型草原植物多样性与初级生产力的影响及途径

王玉冰^1,²,孙毅寒^1,²,丁威³,张恩涛^1,²,李文怀¹,迟永刚^4,^*(),郑淑霞^1,^*()

¹中国科学院植物研究所植被与环境变化国家重点实验室, 北京 100093
²中国科学院大学, 北京 100049
³江西师范大学生命科学学院, 南昌 330022
⁴浙江师范大学地理与环境科学学院, 浙江金华 321004

收稿日期:2019-10-01 修回日期:2020-01-09 出版日期:2020-01-20 发布日期:2020-02-24
通讯作者: 迟永刚,郑淑霞
基金资助:
国家重点研发计划(2016YFC0500801);国家自然科学基金(41671046);国家自然科学基金(31400393);国家自然科学基金(31600365)

Effects and pathways of long-term nitrogen addition on plant diversity and primary productivity in a typical steppe

WANG Yu-Bing^1,²,SUN Yi-Han^1,²,DING Wei³,ZHANG En-Tao^1,²,LI Wen-Huai¹,CHI Yong-Gang^4,^*(),ZHENG Shu-Xia^1,^*()

¹State Key Laboratory of Vegetation and Environmental Change, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China
²University of Chinese Academy of Sciences, Beijing 100049, China
³College of Life Sciences, Jiangxi Normal University, Nanchang 330022, China
⁴College of Geography and Environmental Sciences, Zhejiang Normal University, Jinhua, Zhejiang 321004, China

Received:2019-10-01 Revised:2020-01-09 Online:2020-01-20 Published:2020-02-24
Contact: CHI Yong-Gang,ZHENG Shu-Xia
Supported by:
National Key R&D Program of China(2016YFC0500801);National Natural Science Foundation of China(41671046);National Natural Science Foundation of China(31400393);National Natural Science Foundation of China(31600365)

摘要/Abstract

摘要：

氮(N)沉降对陆地生态系统的结构和功能已产生了重要的影响, N也是中国北方草原植物生长和初级生产力的主要限制性元素。物种多样性和功能多样性是揭示生物多样性对生态系统功能维持机制的关键指标, 然而, 关于长期N添加下草原物种多样性与功能多样性的关系, 及其对初级生产力的影响途径及机制, 尚不十分清楚。为此, 该研究依托在内蒙古典型草原建立的长期N添加实验平台, 实验处理包括1个完全对照(不添加任何肥料)和6个N添加水平(0、1.75、5.25、10.50、17.50和28.00 g·m^-2·a^-1), 研究了长期N添加对典型草原物种多样性、功能多样性和初级生产力的影响大小及途径。结果表明: 1) N添加显著降低了典型草原的物种丰富度和Shannon-Wiener指数, 但对功能多样性(包括功能性状多样性指数和群落加权性状值)无显著的影响。2)结构方程模型分析表明, 功能多样性主要受物种丰富度的影响, 但是物种多样性减少并没有导致功能多样性降低, 其原因主要是功能群组成发生了改变, 即群落内多年生根茎禾草所占比例显著增加, 以致群落加权性状值变化不大。3) N通过影响物种丰富度和功能群组成, 间接影响群落加权性状值, 进而影响群落净初级生产力。其中, 群落加权性状值是最重要的影响因子, 可解释48%的初级生产力变化, 表明初级生产力主要是由群落内优势物种的生物量及功能性状所决定, 因此该研究的结果很好地支持了质量比假说。

关键词: 植物功能性状, 物种多样性, 功能多样性, 群落加权性状, 功能群组成, 生物多样性, 地上净初级生产力

Abstract:

Aims Nitrogen (N) deposition has made great impacts on the structure and function of terrestrial ecosystems in recent decades, and N is known as the main limiting element of plant growth and primary productivity in the northern grassland of China. Species diversity and functional diversity have been widely recognized as key indicators of the mechanisms of biodiversity maintenance for ecosystem functions. However, the effect of long-term N addition on the relationship between species diversity and functional diversity, and the corresponding impacts on primary productivity have rarely been studied; particularly the underpinning mechanisms remain unclear. Our objective is to examine the effects and pathways of long-term N addition on species diversity, functional diversity and primary productivity in a typical steppe.
Methods A manipulative N addition experiment located in a typical steppe of the Nei Mongol grassland has been conducted for 18 years. The experimental design included seven levels of N addition rate (i.e., control, 0, 1.75, 5.25, 10.50, 17.50, 28.00 g·m^-2·a^-1) with nine replicates for each treatment. The plant functional traits of dominant species, species richness and composition, and aboveground net primary productivity (ANPP) were determined. In addition, species diversity, functional attribute diversity and community-weighted mean traits were calculated.
Important findings 1) N addition significantly reduced species richness and Shannon-Wiener index, but had little effect on functional diversity, including functional attribute diversity and community-weighted mean traits. 2) Structural equation model analyses showed that functional diversity was mainly affected by species richness, whereas the decrease in species diversity didn’t lead to the decrease in functional diversity. The community- weighted mean traits did not change under N addition, which was mainly due to the shift in functional group composition, that is, the proportion of perennial rhizome grass in plant community increased greatly under N addition. 3) The effect of N addition on ANPP was through direct pathways of species richness and shift in functional group composition, and further via an indirect pathway of community-weighted mean traits. Particularly, the community-weighted mean traits were the most important factors and explained 48% of total variation in ANPP, implying that the primary productivity is mainly determined by the biomass and functional traits of dominant species in the plant community. Our results well supported the mass-ratio hypothesis.

Key words: plant functional trait, species diversity, functional diversity, community-weighted mean traits, functional group composition, biodiversity, aboveground net primary productivity

王玉冰,孙毅寒,丁威,张恩涛,李文怀,迟永刚,郑淑霞. 长期氮添加对典型草原植物多样性与初级生产力的影响及途径. 植物生态学报, 2020, 44(1): 22-32. DOI: 10.17521/cjpe.2019.0260

WANG Yu-Bing,SUN Yi-Han,DING Wei,ZHANG En-Tao,LI Wen-Huai,CHI Yong-Gang,ZHENG Shu-Xia. Effects and pathways of long-term nitrogen addition on plant diversity and primary productivity in a typical steppe. Chinese Journal of Plant Ecology, 2020, 44(1): 22-32. DOI: 10.17521/cjpe.2019.0260

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图/表 6

图1 氮添加对典型草原物种丰富度(A)和Shannon-Wiener指数(B)的影响(平均值+标准误差)。Control, 完全对照, 不添加任何肥料。不同小写字母表示处理间差异显著(p < 0.05)。

Fig. 1 Effects of N addition on species richness (A) and Shannon-Wiener index (B) in a typical steppe (mean + SE). Control, absolute control, no fertilizer added. Different lowercase letters indicate significant difference among treatments (p < 0.05).

图2 氮添加对典型草原群落功能性状多样性(FAD)的影响(平均值+标准误差)。A 株丛性状多样性, 以株高、株丛生物量和茎叶比计算。B 叶片性状多样性, 以比叶面积、叶片干物质含量和叶片N含量计算。C 整株性状多样性, 以株丛性状和叶片性状共6个指标计算。Control, 完全对照, 不添加任何肥料。各处理间差异不显著(p > 0.05)。

Fig. 2 Effect of N addition on functional attribute diversity (FAD) in a typical steppe (mean + SE). A, Plant trait functional diversity, calculated by plant height (PH), individual biomass (PB) and stem:leaf biomass ratio (SLR). B, Leaf trait functional diversity, calculated by specific leaf area (SLA), leaf dry mass content (LDMC), and leaf N content (LNC). C, Whole-plant trait functional diversity, calculated by six indices of plant and leaf traits. Control, absolute control, no fertilizer added. No significant difference among treatments (p > 0.05).

图3 氮添加对典型草原群落加权性状的影响(平均值+标准误差)。群落性状加权值: PHCWM, 株高; PBCWM, 株丛生物量; SLRCWM, 茎叶比; SLACWM, 比叶面积; LDMCCWM, 叶片干物质含量; LNCCWM, 叶片氮含量。Control, 完全对照, 不添加任何肥料。各处理间差异不显著(p > 0.05)。

Fig. 3 Effects of N addition on community-weighted mean traits in a typical steppe (mean + SE). Community-weighted mean traits included community-weighted plant height (PHCWM), individual biomass (PBCWM), stem:leaf biomass ratio (SLRCWM), specific leaf area (SLACWM), leaf dry mass content (LDMCCWM), and leaf N content (LNCCWM). Control, absolute control, no fertilizer added. No significant difference among treatments (p > 0.05).

图4 氮添加对典型草原6个优势物种相对生物量(RAB)的影响(平均值±标准误差)。PR, 多年生根茎禾草; PB, 多年生丛生禾草; PF, 多年生杂类草。Control, 完全对照, 不添加任何肥料。

Fig. 4 Effects of N addition on the relative aboveground biomass (RAB) of six dominant species in a typical steppe (mean ± SE). PR, perennial rhizome grasses; PB, perennial bunch grasses; PF, perennial forbs. Control, absolute control, no fertilizer added.

图5 氮添加对典型草原群落初级生产力(ANPP)的影响(平均值+标准误差)。Control, 完全对照, 不添加任何肥料。各处理间差异不显著(p > 0.05)

Fig. 5 Effect of N addition on aboveground net primary productivity (ANPP) in a typical steppe (mean + SE). Control, absolute control, no fertilizer added. No significant difference among treatments (p > 0.05).

图6 氮添加对典型草原净初级生产力(ANPP)影响途径的结构方程模型(SEM)分析结果。c2 = 3.611, p = 0.89, df = 8, n = 13, RMSEA < 0.001, AIC = 29.61, CFI = 1.000。实线和虚线分别表示变量之间作用关系显著(p < 0.05)和不显著(p > 0.05)。正负数值表示标准化回归系数(β); *, p < 0.05; **, p < 0.01; ***, p < 0.001; r2值表示某一变量被其他变量的方差解释量。

Fig. 6 Structural equation model (SEM) analyses of pathways of N addition on aboveground net primary productivity (ANPP). Results of SEM fitting: c2 = 3.611, p = 0.89, df = 8, n = 13, RMSEA < 0.001, AIC = 29.61, CFI = 1.000. ANPP, aboveground net primary productivity; CWM, community-weighted mean traits; FAD, functional attribute diversity; PR/PB, biomass ratio of perennial rhizome grass to perennial bunchgrass, indicating a shift in functional group composition. Solid and dashed arrows indicate significant (p < 0.05) and non-significant (p > 0.05) effects, respectively. Values associated with solid arrows represent standardized path coefficients (β); *, p < 0.05; **, p < 0.01; ***, p < 0.001. r2 values associated with response variables indicate the proportion of variation explained by relationships with other variables.

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长期氮添加对典型草原植物多样性与初级生产力的影响及途径

Effects and pathways of long-term nitrogen addition on plant diversity and primary productivity in a typical steppe

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