内蒙古羊草草原不同退化阶段土壤养分与植物功能性状的关系

doi:10.17521/cjpe.2015.0465

摘要/Abstract

摘要：

深入认识植物功能性状的生态学含义, 对于阐明不同自然与人为干扰环境下的群落构建途径, 进一步揭示生态系统服务维持机制具有重要的理论意义。该文以内蒙古锡林河流域羊草(Leymus chinensis)草原不同退化演替阶段的群落为研究对象, 分析了土壤养分与植物功能性状的变化特征及两者之间的关系。结果表明: (1)退化导致土壤养分含量逐渐减少, 全氮和全磷在未退化的羊草+杂类草群落与严重退化的羊草+冷蒿(Artemisia frigida)群落之间差异显著; (2)随着退化演替的进程, 群落最大高度和叶片碳氮比减小, 群落最大高度在未退化的羊草+杂类草群落与轻度退化的羊草+针茅(Stipa sp.)群落之间差异显著, 碳氮比在未退化的羊草+杂类草群落与严重退化的羊草+冷蒿群落之间差异显著; (3)不同退化演替阶段的群落, 其土壤养分对植物功能性状的影响有所差异。在羊草+针茅群落, 速效氮与群落最大高度、叶片木质素含量和叶片碳氮比均呈显著负相关关系。而在羊草+糙隐子草(Cleistogenes squarrosa)群落中, 上述3种植物功能性状则表现为均与全磷含量显著正相关; (4)群落植物功能性状之间的关系也因退化阶段不同而有所不同。在轻度退化的羊草+针茅群落中, 叶片木质素含量与其他4个功能性状显著正相关, 叶片碳氮比与群落最大高度、叶干物质含量、木质素含量呈显著正相关关系。在严重退化的羊草+冷蒿群落中, 所有性状均呈极显著正相关关系。表明植物通过功能性状的协调或组合, 以适应贫瘠的土壤环境。上述结果深化了对典型草原退化演替的认识, 对退化草地的恢复与保护具有一定的指导意义。

关键词: 植物功能性状, 土壤养分, 退化演替, 羊草草原, 锡林河流域

Abstract:

Aims Understanding ecological implications of plant functional traits is helpful in exploring community assembly under different environments of nature and human disturbances, and then to reveal the maintenance mechanism of the ecosystem services. By analyzing vegetation and soil data derived from field observations in Leymus chinensis steppe of Xilin River Basin in Nei Mongol, we aimed to explore the responses of plant functional traits to changing soil nutrients at different degradation stages. Methods We observed 69 plots for both plant community structure and soil attributes using quadrat and soil-drilling methods. Five plant functional traits, namely the specific leaf area (SLA), leaf dry matter content (LDMC), leaf carbon to nitrogen ratio (C:N), leaf lignin content (LLC), and maximum height (MH), were measured for each plot. We also tested soil attributes, such as total nitrogen (TN), total phosphorus (TP), available nitrogen (AN), available phosphorus (AP), and organic carbon (OC). The sixty-nine communities were classified into four groups (undegraded L. chinensis + forbs, slightly degraded L. chinensis + Stipa sp., moderately degraded L. chinensis + Cleistogenes squarrosa, and heavily degraded L. chinensis + Artemisia frigida) using TWINSPAN software. The relationships between plant functional traits and soil nutrient variables were analyzed for the four community groups using the Pearson’s correlation test with SPSS 21.0 software. Important findings (1) The soil nutrients decreased with the grassland degradation process and there were significant differences in TN and TP between the undegraded L. chinensis + forbs and heavily degraded L. chinensis + A. frigida communities; (2) plant functional traits also showed strong differences between the degradation stages. MH and C:N decreased with degradation. A significant difference was observed in MH between the undegraded L. chinensis + forbs and slightly degraded L. chinensis + Stipa sp. communities. The difference in C:N was also significant between the undegraded L. chinensis + forbs and heavily degraded L. chinensis + A. frigida communities; (3) the effects of soil nutrients on plant functional traits changed with grassland degradation. AN was negatively correlated with MH, LLC, and C:N in the slightly degraded L. chinensis + Stipa sp. community. In the moderately degraded L. chinensis + C. squarrosa community, those three traits mentioned above showed significantly positive correlations with TP; (4) while analyzing the degraded grassland, different relationships between plant functional traits were found. In the slightly degraded L. chinensis + Stipa sp. community, LLC was positively correlated with all other traits. Moreover, positive correlations also occurred between C:N and MH, C:N and LDMC, and C:N and LLC. In the heavily degraded L. chinensis + A. frigida community, all traits demonstrated the most significantly positive correlations.

Key words: plant functional traits, soil nutrient, degradation succession, the Leymus chinensis steppe, Xilin River Basin

李丹, 康萨如拉, 赵梦颖, 张庆, 任海娟, 任婧, 周俊梅, 王珍, 吴仁吉, 牛建明. 内蒙古羊草草原不同退化阶段土壤养分与植物功能性状的关系. 植物生态学报, 2016, 40(10): 991-1002. DOI: 10.17521/cjpe.2015.0465

Dan LI, Saruul KANG, Meng-Ying ZHAO, Qing ZHANG, Hai-Juan REN, Jing REN, Jun-Mei ZHOU, Zhen WANG, Ren-Ji WU, Jian-Ming NIU. Relationships between soil nutrients and plant functional traits in different degradation stages of Leymus chinensis steppe in Nei Mongol, China. Chinese Journal of Plant Ecology, 2016, 40(10): 991-1002. DOI: 10.17521/cjpe.2015.0465

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2015.0465

https://www.plant-ecology.com/CN/Y2016/V40/I10/991

图/表 6

图1 样地分布图。

Fig. 1 The distribution map of observation sites.

表1 土壤养分含量与植物功能性状之间的关系

Table 1 Relationships between soil nutrient contents and plant functional traits

类型 Type	指标 Index	全氮 TN	速效氮 AN	全磷 TP	速效磷 AP	有机碳 OC
羊草+杂类草 Leymus chinensis + forbs	SLA	-0.074	-0.208	0.128	-0.247	0.007
	MH	-0.118	0.098	0.031	0.048	-0.167
	LDMC	0.225	0.297	0.208	0.090	0.244
	LLC	-0.361	-0.196	-0.218	-0.316	-0.268
	C:N	-0.275	-0.571^**	-0.043	-0.534^*	-0.126
羊草+针茅 Leymus chinensis + Stipa sp.	SLA	-0.051	-0.305	0.106	0.147	0.019
	MH	-0.398	-0.515^*	-0.037	-0.214	-0.395
	LDMC	0.061	-0.349	0.204	0.142	0.054
	LLC	-0.384	-0.667^**	-0.102	-0.126	-0.382
	C:N	-0.203	-0.697^**	0.068	-0.114	-0.208
羊草+糙隐子草 Leymus chinensis + Cleistogenes squarrosa	SLA	0.052	-0.078	0.034	-0.103	0.144
	MH	0.371	0.210	0.477^*	-0.096	0.222
	LDMC	0.306	-0.257	0.376	0.293	0.049
	LLC	0.509^*	-0.066	0.514^*	0.181	0.286
	C:N	0.223	-0.362	0.562^**	0.262	-0.029
羊草+冷蒿 Leymus chinensis + Artemisia frigida	SLA	0.519	0.127	0.309	0.280	0.416
	MH	0.432	0.140	0.236	0.254	0.332
	LDMC	0.404	0.305	0.202	0.159	0.329
	LLC	0.354	0.144	0.132	0.143	0.250
	C:N	0.608	0.342	0.441	0.260	0.526

图2 不同退化阶段土壤养分含量比较(平均值±标准误差)。I, 羊草+杂类草; II, 羊草+针茅; III, 羊草+糙隐子草; IV, 羊草+冷蒿。ns, 群落间差异不显著。每个子图中不同字母表示不同演替阶段之间差异显著(p < 0.05)。

Fig. 2 Comparison of soil nutrient among different degradation stages (mean ± SE). I, Leymus chinensis + forbs; II, Leymus chinensis + Stipa sp.; III, Leymus chinensis + Cleistogenes squarrosa; IV, Leymus chinensis + Artemisia frigida. ns refers to no significant difference at p > 0.05. Letters in each subgraph indicate significant differences between stages (p < 0.05). AN, available nitrogen; AP, available phosphorus; OC, organic carbon; TN, total nitrogen; TP, total phosphorus.

图3 不同退化阶段植物功能性状比较(平均值±标准误差)。I, 羊草+杂类草; II, 羊草+针茅; III, 羊草+糙隐子草; IV, 羊草+冷蒿。ns, 群落间差异不显著。每个子图中不同字母表示不同演替阶段之间差异显著(p < 0.05)。

Fig. 3 Comparison of plant functional traits among different degradation stages (mean ± SE). I, Leymus chinensis + forbs; II, Leymus chinensis + Stipa sp.; III, Leymus chinensis + Cleistogenes squarrosa; IV, Leymus chinensis + Artemisia frigida. ns refers to no significant difference at p > 0.05. Letters in each subgraph indicate significant differences between stages (p < 0.05). C:N, leaf carbon and nitrogen ratio; LDMC, leaf dry matter content; LLC, leaf lignin content; MH, maximum height; SLA, specific leaf area.

表2 植物功能性状之间的关系

Table 2 Relationships among plant functional traits

类型 Type	指标 Index	比叶面积 SLA	最大高度 MH	叶片干物质含量 LDMC	叶片木质素含量 LLC	叶片碳氮比 C:N
羊草+杂类草 Leymus chinensis + forbs	SLA	1	0.015	-0.435^*	-0.050	0.268
	MH		1	-0.005	0.527^*	0.309
	LDMC			1	0.021	-0.102
	LLC				1	0.575^**
	C:N					1
羊草+针茅 Leymus chinensis + Stipa sp.	SLA	1	0.434	0.425	0.603^**	0.406
	MH		1	0.518^*	0.681^**	0.565^*
	LDMC			1	0.493^*	0.634^**
	LLC				1	0.693^**
	C:N					1
羊草+糙隐子草 Leymus chinensis + Cleistogenes squarrosa	SLA	1	-0.128	-0.080	0.088	0.184
	MH		1	0.330	0.520^*	0.232
	LDMC			1	0.735^**	0.356
	LLC				1	0.388
	C:N					1
羊草+冷蒿 Leymus chinensis + Artemisia frigida	SLA	1	0.961^**	0.838^**	0.935^**	0.944^**
	MH		1	0.854^**	0.919^**	0.892^**
	LDMC			1	0.953^**	0.842^**
	LLC				1	0.906^**
	C:N					1

图4 附录I TWINSPAN群落分类结果^顶端两行数字纵列代表样地编号, 从左至右如: 19、61、25至6样地; 每一行的左侧(前11个字符, 含空格)代表植物种编号, 其中Var是”变量”的缩写; 右侧0、1组合代表物种的分类;表格中央部分的数字与短线表示物种在样地中的重要性, 5代表最重要, 短线代表可以忽略; 表格底部的6行是样地分类结果, 0、1代表不同类群, 首先将样地划分为两类(倒数第6行), 接下来(倒数第5行)是对上述两个类群的再次划分(二分法), 形成4个类群, 以此类推。

Fig.4 Appendix I The result of TWINSPAN classification^ Site numbers are shown in vertical sequence (top-down) by the first two lines on the top of the table. The first three sites coded 19, 61, and 25, for example, are at the very beginning on the left, and the last one is site 6 on the right end. Code of plant species is shown on the very left (the first 11 chars, including blanks) for each line. Var is the abbreviation of “variable”. A group of 0 and 1 on the right demonstrates the classification of plant species; Numbers and dashes appeared in the central part of the table indicate the value of species in sites. The number 5 refers to the most important, and the sign of dash illustrates that the value of species in sites can be neglected; Results of site classification are summarized by the last 6 lines at the bottom of the table, 0 and 1 refer to differentiation. At first, all sites are categorized to 2 groups shown in the line 6 from the end. And then, these two groups are classified further into 4 groups (in the line 5 from the end) by means of dichotomy, and so forth.

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