凋落物输入变化和氮添加对半干旱草原群落生产力及功能群组成的影响

doi:10.17521/cjpe.2020.0126

植物生态学报 ›› 2020, Vol. 44 ›› Issue (8): 791-806.DOI: 10.17521/cjpe.2020.0126

• 研究论文 • 下一篇

凋落物输入变化和氮添加对半干旱草原群落生产力及功能群组成的影响

嘎玛达尔基¹^,², 杨泽¹^,², 谭星儒¹^,², 王珊珊¹^,², 李伟晶¹^,², 游翠海¹^,², 王彦兵¹^,², 张兵伟¹^,³, 任婷婷¹^,², 陈世苹¹^,²^,^*()

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中山大学有害生物控制与资源利用国家重点实验室, 中山大学生命科学学院, 广州 510275

收稿日期:2020-04-30 接受日期:2020-07-01 出版日期:2020-08-20 发布日期:2020-07-09
通讯作者: 陈世苹
作者简介:* spchen@ibcas.ac.cn
基金资助:
国家自然科学基金(41773084);国家重点研发计划(2017YFA0604801);国家重点研发计划(2016YFC0500103)

Effect of altered litter input and nitrogen addition on ecosystem aboveground primary productivity and plant functional group composition in a semiarid grassland

GAMADAERJI ¹^,², YANG Ze¹^,², TAN Xing-Ru¹^,², WANG Shan-Shan¹^,², LI Wei-Jing¹^,², YOU Cui-Hai¹^,², WANG Yan-Bing¹^,², ZHANG Bing-Wei¹^,³, REN Ting-Ting¹^,², CHEN Shi-Ping¹^,²^,^*()

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
³State Key Laboratory of Biocontrol, School of Life Sciences, Sun Yat-sen University, Guangzhou 510275, China

Received:2020-04-30 Accepted:2020-07-01 Online:2020-08-20 Published:2020-07-09
Contact: CHEN Shi-Ping
Supported by:
National Natural Science Foundation of China(41773084);National Key R&D Program of China(2017YFA0604801);National Key R&D Program of China(2016YFC0500103)

摘要/Abstract

摘要：

不同的草原利用方式(围封、放牧和割草等)随着大气氮沉降的不断加剧, 改变了凋落物输入量。凋落物作为连接地上-地下碳循环过程的关键环节, 对草原生态系统生产力和碳循环过程影响显著。氮是草原生产力的主要限制因子, 凋落物输入量的变化对草原生态系统结构和功能的影响仍缺乏长期实验证据支持。该研究在内蒙古半干旱典型草原建立一个凋落物输入变化和氮添加控制实验平台, 通过连续6年对群落生产力和功能群组成的监测, 研究了凋落物添加与去除和氮添加对半干旱草原群落生产力和功能群组成的影响。研究发现: 1)凋落物输入量增加和氮添加均显著提高了群落生产力, 在对照和氮添加处理下, 凋落物去除处理导致生产力分别降低了8.4%和7.6%, 而凋落物添加处理使生产力分别提高了10.7%和6.3%; 2)不同植物功能群对凋落物输入变化和氮添加的响应存在差异, 导致群落功能群结构发生变化。随着凋落物输入量增加和氮添加, 群落优势功能群多年生禾草(包括多年生丛生禾草和多年生根茎禾草)的生物量显著提高, 对群落生产力的贡献增加, 在群落中的优势地位增强; 而另一优势功能群多年生杂类草生物量对凋落物和氮添加处理均无显著响应, 进而导致在氮添加处理下其对群落生物量的贡献比例显著降低; 3)凋落物输入主要改善土壤水分状况, 而氮添加则主要通过提高土壤养分含量, 促进群落生产力, 并通过影响主要功能群生物量, 导致群落结构发生变化。以上结果表明, 适当的草原管理方式如围封禁牧和降低放牧强度等都能通过增加凋落物的输入来提高草原生产力, 维持生态系统稳定性。而适量的氮等养分添加管理也有助于提高草原生产力, 促进其恢复。

关键词: 地上净初级生产力, 植物功能群, 凋落物输入变化, 氮添加, 半干旱草原

Abstract:

Aims Litter is the major input source of soil organic carbon and nutrients in natural ecosystems and considered as a key link between above- and belowground carbon cycles. Changes in litter input amount have been proven to exert significant impacts on plant productivity, community structure, and therefore ecosystem function. In Nei Mongol semiarid grasslands, different grassland management practices such as grazing, clipping, and fencing have caused dramatic changes in litter production and input. In addition, as a nitrogen-limited ecosystem, Nei Mongol semiarid grasslands also experienced increasing nitrogen deposition. However, how do changes in litter input and nitrogen addition impact the community productivity and composition of plant functional groups are still unclear in the semiarid grasslands. In this study, our objectives are: 1) to investigate the effects of altered litter input and nitrogen addition on community productivity; 2) to study the changes in aboveground biomass of different plant functional groups and their contribution to community productivity under different litter input and N addition treatments.
Methods We established a manipulative experiment with altered litter input and nitrogen addition treatments in a semiarid typical grassland in West Ujimqin Banner, Nei Mongol. A randomized block split-plot design was applied with five blocks. Three litter input treatments, including litter removal (C0), control (C1) and litter addition (C2), were assigned randomly in each block. Each plot (6 m × 7 m) of litter input treatment was separated into two subplots. One of the subplots was assigned as the N addition treatment (N1) and another subplot was considered as the control treatment without N addition (N0). In N addition treatment, 15 g N·m^-2·a^-1 N fertilizer (as NH₄NO₃) was applied every year since 2013. Aboveground net primary productivity (ANPP) in community and plant functional group levels of each treatment were determined during the peak season from 2013 to 2018.
Important findings Based on 6-year measurements, we found the following results. 1) Litter input increase and nitrogen addition increased community ANPP. Compared with the control, litter removal treatment significantly decreased ANPP by 8.4% and 7.6% in plots without and with N addition, respectively. Litter addition increased ANPP by 10.7% and 6.3% in plots without and with N addition, respectively. 2) Different responses of plant functional groups to altered litter input and N addition led to a change in plant functional group composition. The biomass of perennial bunch grass (PB) and perennial rhizome grass (PR) increased significantly with the increment of litter and nitrogen, which enhanced their dominant status in the community. 3) Improved soil water condition by litter input and nutrient supply by N addition are the major pathways that enhanced ANPP and changed the functional group composition. These results show that proper management, such as grazing exclusion and reducing grazing intensity, can promote productivity by increasing inputs of litter in semiarid grasslands, which leads to the maintenance of ecosystem stability. Suitable nutrients management, like nitrogen addition, is also helpful for productivity improvement and the recovery of degraded grasslands.

Key words: aboveground net primary productivity (ANPP), plant functional group, altered litter input, nitrogen addition, semiarid grassland

嘎玛达尔基, 杨泽, 谭星儒, 王珊珊, 李伟晶, 游翠海, 王彦兵, 张兵伟, 任婷婷, 陈世苹. 凋落物输入变化和氮添加对半干旱草原群落生产力及功能群组成的影响. 植物生态学报, 2020, 44(8): 791-806. DOI: 10.17521/cjpe.2020.0126

GAMADAERJI , YANG Ze, TAN Xing-Ru, WANG Shan-Shan, LI Wei-Jing, YOU Cui-Hai, WANG Yan-Bing, ZHANG Bing-Wei, REN Ting-Ting, CHEN Shi-Ping. Effect of altered litter input and nitrogen addition on ecosystem aboveground primary productivity and plant functional group composition in a semiarid grassland. Chinese Journal of Plant Ecology, 2020, 44(8): 791-806. DOI: 10.17521/cjpe.2020.0126

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2020.0126

https://www.plant-ecology.com/CN/Y2020/V44/I8/791

图/表 11

表1 2013-2018年凋落物添加处理中每年的凋落物添加量(g·m-2)

Table 1 Annual litter addition amount of litter addition treatment in 2013- 2018 (g·m-2)

处理 Treatment	2013	2014	2015	2016	2017	2018
N0	433 ± 13	199 ± 24	203 ± 26	135 ± 38	101 ± 8	129 ± 16
N1	433 ± 13	250 ± 23	291 ± 50	225 ± 47	158 ± 10	190 ± 17

图1 长期平均(1955-2011年)和实验年份(2012-2018年)生长季(5-9月)降水量及日平均气温。

Fig. 1 Changes in growing season precipitation and daily mean air temperature during the studied period (2012-2018) and the longterm mean growing season precipitation during 1955-2011.

表2 凋落物输入变化(C)和氮添加(N)对土壤微环境影响的重复测量方差分析(RANOVA)结果

Table 2 Effects of altered litter input (C) and nitrogen addition (N) on soil microenvironment evaluated using repeated measures analysis of variance (RANOVA)

	df	DOC含量 DOC content	DON含量 DON content	df	Ts	SWC
区组 Block	4	0.86	0.39	4	15.61^***	3.99^*
凋落物管理 C	2	0.76	0.77	2	14.44^***	10.49^***
氮添加 N	1	28.13^***	61.81^***	1	10.52^**	0.12
年 Year (Y)	2	20.14^***	15.33^***	5	780.49^***	302.17^***
C × N	2	0.06	0.36	2	0.09	0.52
C × Y	4	1.19	4.08^**	10	1.89^#	11.12^***
N × Y	2	8.41^**	5.18^*	5	2.20^#	8.42^***
C × N × Y	4	0.48	3.18^*	10	0.32	4.27^***

图2 2013-2018年不同凋落物输入变化和氮添加处理下, 表层(0-10 cm)土壤温度(Ts)、土壤含水量(SWC)和土壤可溶性有机碳氮(DOC和DON)含量的年际变异(A、C、E、G)和6年平均值(B、D、F、H)(平均值±标准误差)。图中给出了凋落物输入变化(C)和氮添加(N)对生产力影响的差异显著性分析结果(ns, p > 0.1; #, p < 0.1; *, p < 0.05; **, p < 0.01; ***, p < 0.001)。柱状图中的字母表示不同凋落物输入变化和氮添加处理间各指标的多重比较结果, 不同小写字母表示不同凋落物输入变化处理间的差异显著(p < 0.05), 不同大写字母代表不同氮添加处理间的差异显著(p < 0.05)。C0, 凋落物去除; C1, 凋落物对照; C2, 凋落物添加; N0, 无氮添加; N1, 氮添加。

Fig. 2 Inter-annual variations (A, C, E, G) and the 6-year average (B, D, F, H) of soil temperature (Ts), soil water content (SWC), dissolved organic carbon (DOC) and nitrogen (DON) content in surface soil layer (0-10 cm) in different litter input and nitrogen addition treatments during 2013-2018 (mean ± SE). Significance levels were presented to show the effects of altered litter input (C) and nitrogen addition (N) treatments and their interaction (C × N) on these parameters (ns, p > 0. 1; #, p < 0.1; *, p < 0.05; **, p < 0.01; ***, p < 0.001). The letters in the bar graph indicate the results of multiple comparisons among different litter input and nitrogen addition treatments. Different lowercase letters indicate significant difference among three altered litter input treatments (p < 0.05), and uppercase letters indicate the significant difference between two nitrogen treatments (p < 0.05). C0, litter removal; C1, control; C2, litter addition; N0, without nitrogen addition; N1, nitrogen addition.

图3 2013-2018年不同凋落物输入变化和氮添加处理下, 群落地上净初级生产力(ANPP)的年际变异(A)和6年平均值(B)。图中数据均为平均值±标准误差。图中给出了凋落物输入变化(C)和氮添加(N)对生产力影响的差异显著性分析结果(ns, p > 0.1; #, p < 0.1; *, p < 0.05; **, p < 0.01; ***, p < 0.001)。图中不同大写字母表示氮添加对生产力影响的差异显著(p < 0.05)。C0, 凋落物去除; C1, 凋落物对照; C2, 凋落物添加; N0, 无氮添加; N1, 氮添加。

Fig. 3 Inter-annual variation (A) and the 6-year average (B) of aboveground net primary productivity (ANPP) in different litter input and nitrogen addition treatments during 2013-2018. The data in the figure are mean ± SE. Significance levels were presented to show the effects of altered litter input (C) and nitrogen addition (N) treatments and their interaction (C × N) on ANPP (ns, p > 0.1; #, p < 0.1; *, p < 0.05; **, p < 0.01; ***, p < 0.001). Different uppercase letters in the bar graph indicate significant difference results of multiple comparisons between two nitrogen treatments (p < 0.05). C0, litter removal; C1, control; C2, litter addition; N0, without nitrogen addition; N1, nitrogen addition.

表3 凋落物输入变化(C)和氮添加(N)对群落地上净初级生产力(ANPP)及各功能群绝对和相对生物量影响的重复测量方差分析(RANOVA)结果

Table 3 Effects of altered litter input (C) and nitrogen addition (N) on community above-ground net primary productivity (ANPP) and absolute and relative biomass of each functional group evaluated using repeated measures analysis of variance (RANOVA)

	df	地上净初级生产力 ANPP	绝对生物量 Absolute biomass					相对生物量 Relative biomass
	df	地上净初级生产力 ANPP	PB	PR	PF	SS	AB	PB%	PR%	PF%	SS%	AB%
区组Block	4	1.72	1.60	0.66	1.17	1.39	1.16	0.87	0.31	1.11	1.94	0.42
C	2	3.20^#	0.77	3.30^#	0.01	0.66	3.13^#	0.18	1.65	0.36	0.70	3.70^*
N	1	62.87^***	31.22^***	7.60^*	1.45	0.17	42.97^***	9.04^**	0.24	12.50^**	2.81	37.54^***
年Year (Y)	5	57.28^***	15.66^***	22.04^***	17.54^***	1.51	4.32^**	4.87^***	8.73^***	6.41^***	1.50	4.32^**
C × N	2	0.02	0.02	0.02	0.08	1.33	2.65^#	0.08	0.18	0.23	1.21	2.96^#
C × Y	10	0.59	1.31	6.36^***	0.76	0.68	1.85^#	2.78^**	5.03^***	0.86	0.44	1.78^#
N × Y	5	7.64^***	8.05^***	2.63^*	5.27^***	2.46^*	3.91^**	8.59^***	0.30	10.38^***	2.72^*	3.67^**
C × N × Y	10	0.84	0.95	1.63	1.22	0.65	1.81^#	0.39	2.26^*	1.46	0.84	1.70^#

图4 2013-2018年不同凋落物输入变化和氮添加处理下, 各功能群地上净初级生产力(ANPP)年际变异(A、C、E、G、I)和6年平均值(B、D、F、H、J)。图中数据均为平均值±标准误差。图中给出了每个年份以及多年平均凋落物输入变化(C)氮添加(N)及其交互作用(C × N)对各功能群地上净初级生产力影响的差异显著性分析结果(ns, p > 0.1; #, p < 0.1; *, p < 0.05; **, p < 0.01; ***, p < 0.001)。不同大写字母表示不同氮添加处理(N)对各功能群地上净初级生产力影响的多重比较结果差异显著(p < 0.05)。C0, 凋落物去除; C1, 凋落物对照; C2, 凋落物添加; N0, 无氮添加; N1, 氮添加。AB, 一二年生植物; PB, 多年生丛生禾草; PF, 多年生杂类草; PR, 多年生根茎禾草; SS, 灌木与半灌木。

Fig. 4 Inter-annual variations (A, C, E, G, I) and 6-year average (B, D, F, H, J) of aboveground net primary productivity (ANPP) of five functional groups in different litter input and nitrogen addition treatments during 2013-2018. The data in the figure are mean ± SE. Significance levels were presented to show the effects of altered litter input (C) and nitrogen addition (N) treatments and their interaction (C × N) on ANPP of each functional group (ns, p > 0.1; #, p < 0.1; *, p < 0.05; **, p < 0.01; ***, p < 0.001). C0, litter removal; C1, control; C2, litter addition; N0, without nitrogen addition; N1, nitrogen addition. Different uppercase letters in the bar graph indicate significant difference of multiple comparisons between two nitrogen addition treatments. AB, annual and biannual plants; PB, perennial bunch grasses; PF, perennial forbs; PR, perennial rhizome grasses; SS, shrubs and semi-shrubs.

图5 不同凋落物输入变化和氮添加处理下, 各功能群相对生物量多年平均值的变化。图中数据均为平均值±标准误差。星号表示不同氮处理间的差异显著(**, p < 0.01; ***, p < 0.001)。C0, 凋落物移除; C1, 对照; C2, 凋落物添加; N0, 无氮添加; N1, 氮添加。AB, 一二年生植物; PB, 多年生丛生禾草; PF, 多年生杂类草; PR, 多年生根茎禾草; SS, 灌木与半灌木。

Fig. 5 Variations in the relative biomass of the five functional groups in different litter input and nitrogen addition treatments. The data in the figure are mean ± SE. Asterisk indicated significant difference between different nitrogen treatments (**, p < 0.01; ***, p < 0.001). C0, litter removal; C1, control; and C2, litter addition; N0, nitrogen-free control; N1, nitrogen addition. AB, annual and biannual plants; PB, perennial bunch grasses; PF, perennial forbs; PR, perennial rhizome grasses; SS, shrubs, and semi-shrubs.

图6 不同氮处理条件下, 地上净初级生产力(ANPP)、土壤温度(Ts)、土壤含水量(SWC)及其季节变异(CVSWC)与上一年添加的凋落物质量间的相关关系。图中标出相关系数(R2)和差异显著性分析结果(*, p < 0.05; **, p < 0.01; ***, p < 0.001)。N0, 无氮添加; N1, 氮添加。

Fig. 6 Correlation analysis of aboveground net primary productivity (ANPP), soil temperature (Ts), soil water content (SWC) and its variation (CVSWC) with litter biomass added in the previous year under each nitrogen treatment. The correlation coefficient (R2) and significance levels are presented in the figure (*, p < 0.05; **, p < 0.01; ***, p < 0.001). N0, nitrogen-free control; N1, nitrogen addition.

图7 群落地上净初级生产力与生长季降水量(A)、土壤温度(B)、土壤含水量(C)和土壤含水量季节变异(D)之间的关系。图中标出相关系数(R2)和差异显著性分析结果(ns, p > 0.05; *, p < 0.05; **, p < 0.01; ***, p < 0.001)。C0, 凋落物去除; C1, 凋落物对照; C2, 凋落物添加; N0, 无氮添加; N1, 氮添加。

Fig. 7 Correlation analysis between aboveground net primary productivity (ANPP) and precipitation (A), soil temperature (Ts)(B), soil water content (SWC)(C), and seasonal variation of soil water content (CVSWC)(D) during the growing season. The correlation coefficient (R2) and the significance levels are presented in the figure (ns, p > 0.05; *, p < 0.05; **, p < 0.01; ***, p < 0.001). C0, litter removal; C1, control; C2, litter addition; N0, without nitrogen addition; N1, nitrogen addition.

图8 凋落物输入变化和氮添加通过土壤环境影响群落功能群组成及地上净初级生产力(ANPP)的结构方程模型。AIC, 赤池信息量; BIC, 贝叶斯信息量; Fisher’s Chi, 费舍尔精确检验值。

Fig. 8 Structural equation model (SEM) on how litter manipulation and nitrogen addition affect the community functional group composition and aboveground net primary productivity (ANPP) through soil environment. CVSWC, variance of soil water content; DOM, soil dissolved organic matter (the first principal component of DOC and DON, which explained 0.88 of them); PB, perennial bunch grasses; PF, perennial forbs; PR, perennial rhizome grasses; SWC, soil water content; Ts, soil temperature. AIC, Akaike Information Criterion; BIC, Bayesian Information Criterion; Fisher’s Chi, result of Fisher exact test.

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凋落物输入变化和氮添加对半干旱草原群落生产力及功能群组成的影响

Effect of altered litter input and nitrogen addition on ecosystem aboveground primary productivity and plant functional group composition in a semiarid grassland

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