海拔对青海湖流域群落水平植物功能性状的影响

doi:10.17521/cjpe.2020.0140

摘要/Abstract

摘要：

海拔变化会引起气压、温度、降水、土壤湿度和风速等环境因子发生急剧变化, 植物功能性状-海拔的相互关系对于预测全球变化背景下山地植物的适应方式具有重要意义。该研究在青海湖流域海拔3 400-4 200 m范围内布设了5个样地(海拔间隔约200 m), 通过植物群落调查, 测定植物功能性状和土壤理化性质, 结合气象数据, 探讨了海拔对青海湖流域群落水平植物功能性状的影响。结果如下: (1)群落加权平均植株高度(H)、叶片干物质含量(LDMC)、叶片碳氮比(C:N)和叶片氮磷比(N:P)随海拔升高显著降低, 比根表面积(SRA)随海拔升高波动下降, 比叶面积(SLA)、叶片氮含量(LNC)和叶片磷含量(LPC)随海拔升高显著升高, 叶片碳含量(LCC)、比根长(SRL)和根组织密度(RTD)随海拔未发生显著变化。(2)所有性状的变异来源以物种组成变化为主, N:P和LPC的种内性状变异与物种组成变化呈现正的协变效应, 其余性状为负的协变效应。(3)降水和0- 10 cm土层土壤养分含量对SLA变化的解释率较高, 温度和10-20 cm土层土壤养分含量对其余性状随海拔变化的解释率较高。以上结果表明青海湖流域植物群落主要通过物种更替来适应随海拔升高而剧烈变化的环境, 且各群落中的非优势种倾向于占据与优势种相反的性状空间来提高资源利用率, 随海拔变化的热量和深层土壤养分含量是群落水平植物功能性状变化的主要影响因子。

关键词: 海拔, 群落水平性状, 叶片化学计量, 种内变异, 物种组成变化

Abstract:

Aims Altitude has prominent effects on many environmental factors, such as atmospheric pressure, temperature, precipitation, soil moisture and wind velocity. The relationship between plant functional traits and altitude are critical for predicting the effects of climate change on montane plants. Our objective is to examine the effect of altitude on community-level plant functional traits in the Qinghai Lake Basin, China.
Methods Five sites were selected with 200 m increase in altitude (3 400-4 200 m) in the Qinghai Lake Basin, China. Community structure, plant functional traits, soil property and atmospheric factors were surveyed and analyzed in this study. Community-weighted mean functional traits (CWM) was calculated according to the relative abundance of species.
Important findings The results showed that: (1) Community-weighted mean plant height (H), leaf dry matter content (LDMC), leaf C:N ratio (C:N) and leaf N:P ratio (N:P) decreased significantly along altitude, while specific root surface area (SRA) fluctuated with altitude. Specific leaf area (SLA), leaf nitrogen content (LNC) and leaf phosphorus content (LPC) increased significantly along altitude, while altitude had no significant effect on leaf carbon content (LCC), root tissue density (RTD) and specific root length (SRL). (2) The variation in CWM along altitude could be explained by species turnover more rather than intraspecific variability. N:P and LPC had a positive covariation, other CWM had a negative covariation. (3) Precipitation and 0-10 cm depth soil nutrients content explained the largest proportion change of SLA. Temperature and 10-20 cm depth soil nutrients content explained the largest proportion change of other CWM along altitude. Overall, these findings suggested that the plant communities in our study adapted to altitude through species turnover, and the non-dominant species tended to occupy opposite trait spaces to the dominant species in the Qinghai Lake Basin, China. Temperature and deeper soil nutrients content had significant effects on CWM along altitude.

Key words: altitude, community-level trait, leaf stoichiometry, intraspecific variability, species turnover

向响, 黄永梅, 杨崇曜, 李泽卿, 陈慧颖, 潘莹萍, 霍佳璇, 任梁. 海拔对青海湖流域群落水平植物功能性状的影响. 植物生态学报, 2021, 45(5): 456-466. DOI: 10.17521/cjpe.2020.0140

XIANG Xiang, HUANG Yong-Mei, YANG Chong-Yao, LI Ze-Qing, CHEN Hui-Ying, PAN Ying-Ping, HUO Jia-Xuan, REN Liang. Effect of altitude on community-level plant functional traits in the Qinghai Lake Basin, China. Chinese Journal of Plant Ecology, 2021, 45(5): 456-466. DOI: 10.17521/cjpe.2020.0140

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2020.0140

https://www.plant-ecology.com/CN/Y2021/V45/I5/456

图/表 8

图1 青海湖流域采样点分布图。

Fig. 1 Distribution of sampling sites in the Qinghai Lake Basin, China.

表1 青海湖流域研究样地的地理位置和植被特征

Table 1 Geographical and vegetation characteristics of each site in the Qinghai Lake Basin, China

序号 Serial number	海拔 Altitude (m)	群系 Alliance	优势种 Dominant species	坡度 Slope (°)	土壤类型 Soil type
1	3 373	紫花针茅+西北针茅草地 Stipa purpurea + S. sareptana Tussock Grassland Aliance	紫花针茅、西北针茅 S. purpurea, S. sareptana	<5	砂质壤土 Sandy loam
2	3 626	紫花针茅草地 S. purpurea Tussock Grassland Aliance	紫花针茅 S. purpurea	11	砂质壤土 Sandy loam
3	3 719	高山嵩草+紫花针茅草地 Kobresia pygmaea + S. purpurea Tussock Grassland Aliance	高山嵩草、紫花针茅 K. pygmaea, S. purpurea	<5	砂质壤土 Sandy loam
4	3 973	高山嵩草草地 K. pygmaea Tussock Grassland Aliance	高山嵩草 K. pygmaea	13	壤质细砂土 Loam fine sand
5	4 222	唐古红景天高山垫状植被 Rhodiola tangutica Alpine Cushion Vegetation	唐古红景天 R. tangutica	<5	砂质壤土 Sandy loam

表2 本研究选取的植物功能性状指标及计算公式

Table 2 Plant functional traits and their equations selected in this study

植物功能性状 Plant functional trait	简写 Abbreviation	单位 Unit	计算公式 Equation
植株高度 Plant height	H	cm	-
比叶面积 Specific leaf area	SLA	cm²·g^-1	叶面积/叶片干质量 Leaf area/dry mass
叶片干物质含量 Leaf dry matter content	LDMC	g·g^-1	叶干质量/叶饱和鲜质量 Leaf dry mass/saturated fresh mass
叶片氮含量 Leaf nitrogen content	LNC	mg·g^-1	-
叶片磷含量 Leaf phosphorus content	LPC	mg·g^-1	-
叶片碳含量 Leaf carbon content	LCC	mg·g^-1	-
叶片碳氮比 Leaf C:N ratio	C:N	-	-
叶片氮磷比 Leaf N:P ratio	N:P	-	-
比根长 Specific root length	SRL	cm·g^-1	细根长度/细根干质量 Fine root length/dry mass
比根表面积 Specific root surface area	SRA	cm²·g^-1	细根表面积/细根干质量 Fine root surface area/dry mass
根组织密度 Root tissue density	RTD	g·cm^-3	细根干质量/细根体积 Fine root dry mass/volume

表3 青海湖流域各样地的气候和土壤营养元素含量特征(平均值±标准误, n = 3)

Table 3 Climate and soil nutrient content characteristics of each site in the Qinghai Lake Basin, China (mean ± SE, n = 3)

序号 Serial number	年平均气温 Mean annual air temperature (℃)	年降水量 Mean annual precipitation (mm)	0-10 cm土壤全氮含量 Soil total nitrogen content in 0-10 cm layer (mg·g^-1)	0-10 cm土壤全磷含量 Soil total phosphorus content in 0-10 cm layer (mg·g^-1)	10-20 cm土壤全氮含量 Soil total nitrogen content in 10-20 cm layer (mg·g^-1)	10-20 cm土壤全磷含量 Soil total phosphorus content in 10-20 cm layer (mg·g^-1)
1	1	348	0.39 ± 0.003^c	0.66 ± 0.164^a	0.34 ± 0.033^b	0.62 ± 0.001^ab
2	-1	434	0.39 ± 0.003^c	0.62 ± 0.008^a	0.31 ± 0.033^c	0.63 ± 0.004^a
3	-2	430	0.43 ± 0.003^b	0.65 ± 0.004^a	0.26 ± 0.000^d	0.57 ± 0.015^bc
4	-4	401	0.49 ± 0.001^a	0.59 ± 0.004^a	0.38 ± 0.000^a	0.59 ± 0.002^b
5	-6	371	0.32 ± 0.003^d	0.53 ± 0.005^a	0.18 ± 0.033^e	0.53 ± 0.007^d

图2 群落加权平均性状随海拔变化的单因素方差分析(平均值±标准误)。不同小写字母表示该性状在不同海拔间具有显著差异(p < 0.05)。C:N, 叶片碳氮比; H, 植株高度; LCC, 叶片碳含量; LDMC, 叶片干物质含量; LNC, 叶片氮含量; LPC, 叶片磷含量; N:P, 叶片氮磷比; RTD, 根组织密度; SLA, 比叶面积; SRA, 比根表面积; SRL, 比根长。

Fig. 2 The one-way ANOVA of community-weighted mean functional traits along altitude (mean ± SE). Different lowercase letters indicate significant differences between different altitudes (p < 0.05). C:N, leaf carbon to nitrogen ratio; H, plant height; LCC, leaf carbon content; LDMC, leaf dry matter content; LNC, leaf nitrogen content; LPC, leaf phosphorus content; N:P, leaf nitrogen to phosphorus ratio; RTD, root tissue density; SLA, specific leaf area; SRA, specific root surface area; SRL, specific root length.

图3 群落加权平均性状变异来源分解。竖线与方框顶部(种内性状变异+物种组成变化)的距离代表协变效应量, 竖线与方框相交时协变效应为负, 不相交时协变效应为正。*, p < 0.05; **, p < 0.01。性状简写见表2。

Fig. 3 Decomposition of total variability in community- weighted mean functional traits. The space between the top of the column and the bar corresponds to the effect of covariation. If the bar is above the column, the covariation is positive, if the bar crosses the column, the covariation is negative. *, p < 0.05; **, p < 0.01. See Table 2 for abbreviation of traits.

图4 六个环境因子的主成分分析。MAP, 年降水量(mm); N10, 0-10 cm土壤全氮含量(mg·g-1); N20, 10-20 cm土壤全氮含量(mg·g-1); P10, 0-10 cm土壤全磷含量(mg·g-1); P20, 10-20 cm土壤全磷含量(mg·g-1); t, 年平均气温(℃)。

Fig. 4 Principal component analysis (PCA) of six environmental factors. MAP, mean annual precipitation (mm); N10, soil total nitrogen content in 0-10 cm layer (mg·g-1); N20, soil total nitrogen content in 10-20 cm layer (mg·g-1); P10, soil total phosphorus content in 0-10 cm layer (mg·g-1); P20, soil total phosphorus content in 10-20 cm layer (mg·g-1); t, mean annual air temperature (°C).

图5 环境因子PC1与PC2对群落加权平均性状变化的解释率。PC1, 热量和深层养分因子; PC2, 降水和表层养分因子。性状简写见表2。

Fig. 5 Proportion of variance in community-weighted mean functional traits explained by PC1 and PC2. PC1, temperature and deep soil nutrient content; PC2, precipitation and surface soil nutrient content. See Table 2 for abbreviation of traits.

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