土壤环境因子对荒漠植物叶性状的驱动机制

doi:10.17520/biods.2025158

生物多样性 ›› 2025, Vol. 33 ›› Issue (11): 25158. DOI: 10.17520/biods.2025158 cstr: 32101.14.biods.2025158

• 研究报告: 植物多样性 • 上一篇下一篇

土壤环境因子对荒漠植物叶性状的驱动机制

琚晓千¹^,²^,³, 田赟¹^,²^,³^,^*(), 徐铭泽⁴^,⁵, 代远萌⁶, 李满乐⁷, 周煜涵¹^,², 刘鹏¹^,²^,³, 贾昕¹^,²^,³, 查天山¹^,²^,³

¹ 北京林业大学水土保持学院, 北京 100083
² 宁夏盐池毛乌素沙地生态系统国家定位观测研究站, 宁夏盐池 751500
³ 北京林业大学水土保持国家林业和草原局重点实验室, 北京 100083
⁴ 自然资源部重庆典型矿区生态修复野外科学观测研究站(重庆地质矿产研究院), 重庆 401120
⁵ 中国农业大学资源与环境学院, 北京 100089
⁶ 北京松山国家级自然保护区管理处, 北京 102115
⁷ 河南省水土保持监测总站, 郑州 450008

收稿日期:2025-04-29 接受日期:2025-09-16 出版日期:2025-11-20 发布日期:2025-12-26
通讯作者: 田赟
基金资助:
国家自然科学基金(31901366);国家自然科学基金(32071842);中央高校基本科研业务费专项资金(FTYX202526);中央高校基本科研业务费专项资金(FTYX202528);中央高校基本科研业务费专项资金(FTYX202533)

How soil environmental factors shape leaf traits in arid-land vegetation

Xiaoqian Ju¹^,²^,³, Yun Tian¹^,²^,³^,^*(), Mingze Xu⁴^,⁵, Yuanmeng Dai⁶, Manle Li⁷, Yuhan Zhou¹^,², Peng Liu¹^,²^,³, Xin Jia¹^,²^,³, Tianshan Zha¹^,²^,³

¹ School of Soil and Water Conservation, Beijing Forestry University, Beijing 100083, China
² Yanchi Ecological Research Station of the Mu Us Sandy Land, Yanchi, Ningxia 751500, China
³ Key Laboratory for Soil and Water Conservation of State Forestry and Grassland Administration, Beijing Forestry University, Beijing 100083, China
⁴ Observation and Reaserch Station of Ecological Restoration for Chongqing Typical Mining Area, Ministry of Natural Resources (Chongqing Institude of Geology and Mineral Resource), Chongqing 401120, China
⁵ College of Resources and Environmental Sciences, China Agricultural University, Beijing 100089, China
⁶ Beijing Songshan National Nature Reserve Administration, Beijing 102115, China
⁷ Soil and Water Conservation Monitoring Gernal Station of Henan Province, Zhengzhou 450008, China

Received:2025-04-29 Accepted:2025-09-16 Online:2025-11-20 Published:2025-12-26
Contact: Yun Tian
Supported by:
National Natural Science Foundation of China(31901366);National Natural Science Foundation of China(32071842);Fundamental Research Funds for the Central Universities(FTYX202526);Fundamental Research Funds for the Central Universities(FTYX202528);Fundamental Research Funds for the Central Universities(FTYX202533)

摘要/Abstract

摘要：

植物性状是连接植物与环境之间的桥梁, 功能群作为研究其关系的基本单元, 承载着响应环境变化的主要信息。研究植物功能群组成和叶性状如何随资源梯度变化, 对于理解荒漠植物的环境适应策略至关重要。本研究以毛乌素沙地黑沙蒿(Artemisia ordosica)群落为研究对象, 选取4个典型恢复阶段(D1: 半固定沙地; D2: 固定沙地; D3: 土壤生物结皮固定沙地; D4: 草本植物固定沙地), 重点分析了不同植物功能群(禾本科、非禾本科和灌木)叶性状与物种组成的阶段性演变规律, 并探讨了影响二者变化的主要土壤环境因子。结果表明: 随着沙地逐步固定, 植物功能群组成趋于复杂化, 其中禾本科和非禾本科种类和数量增加, 而灌木则无明显变化。D1、D2阶段植物功能群叶性状表现为较高的比叶面积和较低的叶干物质含量(leaf dry matter content, LDMC), 表明植物优先投资叶片形态建成以提高资源利用能力; 禾本科具有较高的LDMC和碳同化能力, 适应胁迫环境的能力更强。冗余分析结果表明, 土壤含水量(soil water content, SWC)、土壤碳氮比(soil carbon to nitrogen ratio, SC : SN)和土壤有机碳含量(soil organic carbon content, SOC)对3种植物功能群叶性状影响极显著(P < 0.01)。方差分解和层次分割分析显示, 9个土壤环境因子解释了植物叶性状总变差的18.5%, 其中SC : SN的单独效应最大(4.13%), 且SC : SN和SWC与大多数因子存在共同效应。本研究综合运用多种统计分析方法, 系统识别了荒漠植物性状变异的关键土壤驱动因子及其协同效应, 为区域植被恢复和生态系统可持续管理提供了坚实的理论基础。

关键词: 叶性状, 土壤环境因子, 冗余分析, 固沙阶段, 植物功能群

Abstract

Aims: Plant traits serve as a bridge linking plants and their environment. As the basic unit for studying this relationship, functional groups carry key information on responses to environmental changes. Investigating how the composition of plant functional groups and leaf traits varying along resource gradients is crucial for understanding the environmental adaptation strategies of desert plants.

Methods: This study focused on the Artemisia ordosica communities in the Mu Us sandy land including four typical restoration stages: semi-fixed dune (D1), fixed dune (D2), fixed dune with biological soil crusts (D3), fixed dune with abundant herbaceous plants (D4). Plant communities were categorized into three functional groups: Poaceae, non- Poaceae, and shrub. We revealed the species composition of different plant functional groups at each restoration stage, the characteristics of their leaf traits, and the key soil environmental factors, with redundancy analysis (RDA) and variation partitioning.

Results: As the sand dunes gradually stabilized, the composition of plant functional groups became more complex, and the number of Poaceae and non-Poaceae species showed an increasing trend. However, shrub species showed relatively stable. Leaf traits of plant functional groups during the D1 and D2 stages exhibited higher specific leaf area (SLA) and lower leaf dry matter content (LDMC), suggesting that plants prioritized investment in leaf morphological development to enhance resource utilization under harsh environmental conditions. Among these, grass had higher LDMC and carbon assimilation capacity, which enhanced their ability to adapt to stressful environments. RDA revealed that soil water content (SWC), soil carbon to nitrogen ratio (SC : SN), and soil organic carbon content (SOC) significantly influenced the leaf trait assemblages of all three plant functional groups (P < 0.01). Variation partitioning and hierarchical partitioning analysis indicated that nine soil factors explained 18.5% of the total variance in plant leaf traits. The SC : SN had the largest individual effect (4.13%), and both SC : SN and SWC exhibited co-effects with most other factors.

Conclusion: This study employed multiple statistical analytical approaches to identify the key soil drivers of trait variation in desert plants and their synergistic effects, providing a robust theoretical foundation for regional vegetation restoration and sustainable ecosystem management.

Key words: leaf traits, soil environmental factors, redundancy analysis, sand-fixation stages, plant functional groups

琚晓千, 田赟, 徐铭泽, 代远萌, 李满乐, 周煜涵, 刘鹏, 贾昕, 查天山 (2025) 土壤环境因子对荒漠植物叶性状的驱动机制. 生物多样性, 33, 25158. DOI: 10.17520/biods.2025158.

Xiaoqian Ju, Yun Tian, Mingze Xu, Yuanmeng Dai, Manle Li, Yuhan Zhou, Peng Liu, Xin Jia, Tianshan Zha (2025) How soil environmental factors shape leaf traits in arid-land vegetation. Biodiversity Science, 33, 25158. DOI: 10.17520/biods.2025158.

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链接本文: https://www.biodiversity-science.net/CN/10.17520/biods.2025158

https://www.biodiversity-science.net/CN/Y2025/V33/I11/25158

图/表 7

表1 毛乌素沙地不同固沙阶段样地位置及土壤理化因子变化特征

Table 1 Location information and variation characteristics of soil physicochemical properties across different sand-fixation stages in Mu Us sandy land

固沙阶段 Sand-fixation stages	半固定沙地 Semi-fixed dune (D1)	固定沙地 Fixed dune (D2)	土壤结皮固定沙地 Fixed dune with biological soil crusts (D3)	草本植物固定沙地 Fixed dune with abundant herbaceous plants (D4)
样地图 Plot photograph
经纬度 Latitude and longitude	107.23° N, 37.70° E	107.23° N, 37.71° E	107.23° N, 37.71° E	107.22° N, 37.70° E
海拔 Elevation (m)	1,549	1,541	1,508	1,549
土壤容重 Soil bulk density (BD) (g/cm)	1.54 ± 0.03^a	1.43 ± 0.12^b	1.62 ± 0.03^a	1.57 ± 0.02^a
土壤碳氮比 Soil carbon to nitrogen ratio (SC : SN)	2.53 ± 0.42^c	7.44 ± 1.70^a	3.97 ± 0.65^b	4.84 ± 1.04^b
土壤碳磷比 Soil carbon to phosphorus ratio (SC : SP)	12.63 ± 9.35^a	14.33 ± 4.80^a	13.85 ± 3.90^a	13.41 ± 4.47^a
土壤氮磷比 Soil nitrogen to phosphorus ratio (SN : SP)	4.98 ± 3.63^a	2.02 ± 0.79^b	3.59 ± 1.29^ab	2.95 ± 1.26^ab
土壤有机碳含量 Soil organic carbon content (SOC) (g/kg)	0.75 ± 0.07^c	2.69 ± 1.32^a	1.20 ± 0.27^bc	1.91 ± 0.64^ab
土壤温度 Soil temperature (ST) (℃)	25.18 ± 1.37^a	23.22 ± 2.82^a	24.87 ± 3.06^a	24.47 ± 2.11^a
土壤含水量 Soil water content (SWC) (%)	0.08 ± 0.02^a	0.10 ± 0.03^a	0.03 ± 0.01^b	0.04 ± 0.02^b
土壤全氮含量 Soil total nitrogen content (TN) (g/kg)	0.22 ± 0.03^b	0.31 ± 0.12^a	0.24 ± 0.02^ab	0.30 ± 0.02^ab
土壤全磷含量 Soil total phosphorus content (TP) (g/kg)	0.05 ± 0.02^b	0.19 ± 0.14^a	0.07 ± 0.02^b	0.12 ± 0.06^ab

表2 毛乌素沙地不同固沙阶段不同植物功能群的物种组成

Table 2 Species composition of different plant functional groups across different sand-fixation stages in Mu Us sandy land

固沙阶段 Sand-fixation stages	禾本科 Poaceae	非禾本科 Non-Poaceae	灌木 Shrub
半固定沙地 Semi-fixed dune (D1)	赖草 Leymus secalinus	虫实 Corispermum puberulum 草木樨状黄芪 Astragalus melilotoides 沙蓬 Agriophyllum squarrosum 乳白黄芪 Astragalus galactites	黑沙蒿 Artemisia ordosica 蒙古羊柴 Corethrodendron fruticosum 沙芥 Pugionium cornutum
固定沙地 Fixed dune (D2)	赖草 Leymus secalinus 沙生针茅 Stipa glareosa 隐子草 Cleistogenes caespitosa	虫实 Corispermum puberulum 狗娃花 Aster hispidus 乳白黄芪 Astragalus galactites 草木樨状黄芪 Astragalus melilotoides 砂珍棘豆 Oxytropis racemosa 大翅蓟 Onopordum acanthium	黑沙蒿 Artemisia ordosica 华北白前 Cynanchum komarovii 柠条锦鸡儿 Caragana arborescens 达乌里胡枝子 Lespedeza daurica
土壤结皮固定沙地 Fixed dune with biological soil crusts (D3)	沙生针茅 Stipa glareosa 中华草沙蚕 Tripogon chinensis 隐子草 Cleistogenes caespitosa	虫实 Corispermum puberulum 角蒿 Incarvillea sinensis 乳白黄芪 Astragalus galactites 狗娃花 Aster hispidus 银灰旋花 Convolvulus ammannii	黑沙蒿 Artemisia ordosica 蒙古羊柴 Corethrodendron fruticosum 华北白前 Cynanchum komarovii 达乌里胡枝子 Lespedeza daurica
草本植物固定沙地 Fixed dune with abundant herbaceous plants (D4)	赖草 Leymus secalinus 沙生针茅 Stipa glareosa 隐子草 Cleistogenes caespitosa 沙芦苇 Agropyron mongolicum 中华草沙蚕 Tripogon chinensis	虫实 Corispermum puberulum 角蒿 Incarvillea sinensis 狗娃花 Aster hispidus 甘草 Glycyrrhiza uralensis 大翅蓟 Onopordum acanthium 苦豆子 Sophora alopecuroides 鹅绒藤 Cynanchum chinense 雾冰藜 Grubovia dasyphylla	黑沙蒿 Artemisia ordosica 蒙古羊柴 Corethrodendron fruticosum 华北白前 Cynanchum komarovii

表2 毛乌素沙地不同固沙阶段不同植物功能群的物种组成

Table 2 Species composition of different plant functional groups across different sand-fixation stages in Mu Us sandy land

固沙阶段 Sand-fixation stages	禾本科 Poaceae	非禾本科 Non-Poaceae	灌木 Shrub
半固定沙地 Semi-fixed dune (D1)	赖草 Leymus secalinus	虫实 Corispermum puberulum 草木樨状黄芪 Astragalus melilotoides 沙蓬 Agriophyllum squarrosum 乳白黄芪 Astragalus galactites	黑沙蒿 Artemisia ordosica 蒙古羊柴 Corethrodendron fruticosum 沙芥 Pugionium cornutum
固定沙地 Fixed dune (D2)	赖草 Leymus secalinus 沙生针茅 Stipa glareosa 隐子草 Cleistogenes caespitosa	虫实 Corispermum puberulum 狗娃花 Aster hispidus 乳白黄芪 Astragalus galactites 草木樨状黄芪 Astragalus melilotoides 砂珍棘豆 Oxytropis racemosa 大翅蓟 Onopordum acanthium	黑沙蒿 Artemisia ordosica 华北白前 Cynanchum komarovii 柠条锦鸡儿 Caragana arborescens 达乌里胡枝子 Lespedeza daurica
土壤结皮固定沙地 Fixed dune with biological soil crusts (D3)	沙生针茅 Stipa glareosa 中华草沙蚕 Tripogon chinensis 隐子草 Cleistogenes caespitosa	虫实 Corispermum puberulum 角蒿 Incarvillea sinensis 乳白黄芪 Astragalus galactites 狗娃花 Aster hispidus 银灰旋花 Convolvulus ammannii	黑沙蒿 Artemisia ordosica 蒙古羊柴 Corethrodendron fruticosum 华北白前 Cynanchum komarovii 达乌里胡枝子 Lespedeza daurica
草本植物固定沙地 Fixed dune with abundant herbaceous plants (D4)	赖草 Leymus secalinus 沙生针茅 Stipa glareosa 隐子草 Cleistogenes caespitosa 沙芦苇 Agropyron mongolicum 中华草沙蚕 Tripogon chinensis	虫实 Corispermum puberulum 角蒿 Incarvillea sinensis 狗娃花 Aster hispidus 甘草 Glycyrrhiza uralensis 大翅蓟 Onopordum acanthium 苦豆子 Sophora alopecuroides 鹅绒藤 Cynanchum chinense 雾冰藜 Grubovia dasyphylla	黑沙蒿 Artemisia ordosica 蒙古羊柴 Corethrodendron fruticosum 华北白前 Cynanchum komarovii

表3 毛乌素沙地不同固沙阶段不同植物功能群多样性指数

Table 3 Diversity indices of different plant functional groups across different sand-fixation stages in Mu Us sandy land

固沙阶段 Sand-fixation stages	功能群 Functional groups	相对多度 Relative abundance (%)	相对盖度 Relative coverage (%)	相对频度 Relative frequency (%)	Shannon-Wiener 多样性指数 Shannon-Wiener diversity index (H′)	物种丰富度指数 Species richness index (S)
半固定沙地 Semi-fixed dune (D1)	灌木 Shrub	11	94	48	0.70 ± 0.28^b	5.00 ± 1.00^b
	禾本科 Poaceae	3	1	8
	非禾本科 Non-Poaceae	86	5	44
固定沙地 Fixed dune (D2)	灌木 Shrub	38	79	32	1.86 ± 0.22^a	11.67 ± 0.58^a
	禾本科 Poaceae	34	2	28
	非禾本科 Non-Poaceae	28	19	40
土壤结皮固定沙地 Fixed dune with biological soil crusts (D3)	灌木 Shrub	68	98	46	1.57 ± 0.29^a	8.67 ± 2.08^ab
	禾本科 Poaceae	9	1	20
	非禾本科 Non-Poaceae	23	1	34
草本植物固定沙地 Fixed dune with abundant herbaceous plants (D4)	灌木 Shrub	36	89	30	1.89 ± 0.19^a	12.33 ± 1.15^a
	禾本科 Poaceae	35	8	30
	非禾本科 Non-Poaceae	29	3	40

图1 不同固沙阶段各植物功能群叶性状差异。LA: 叶面积; SLA: 比叶面积; LTD: 叶组织密度; LDMC: 叶干物质含量; LCC: 叶片碳含量; LNC: 叶片氮含量; LPC: 叶片磷含量; LC : LN: 叶片碳氮比; LN : LP: 叶片氮磷比; LC : LP: 叶片碳磷比; D1: 半固定沙地; D2: 固定沙地; D3: 土壤结皮固定沙地; D4: 草本植物固定沙地。不同大写字母表示不同固沙阶段间差异显著, 不同小写字母表示不同功能群间差异显著(P < 0.05)。

Fig. 1 Differences in leaf traits of plant functional groups across different sand-fixation stages. LA, Leaf area; SLA, Specific leaf area; LTD, Leaf tissue density; LDMC, Leaf dry matter content; LCC, Leaf carbon content; LNC, Leaf nitrogen content; LPC, Leaf phosphorus content; LC : LN, Leaf carbon to nitrogen ratio; LN : LP, Leaf nitrogen to phosphorus ratio; LC : LP, Leaf carbon to phosphorus ratio; D1, Semi-fixed dune; D2, Fixed dune; D3, Fixed dune with biological soil crusts; D4, Fixed dune with abundant herbaceous plants. Upper case letters indicate significant differences among different sand-fixation stages and lowercase letters indicate significant differences among different functional groups (P < 0.05).

图2 不同固沙阶段不同植物功能群叶性状集合分布(A-C)及叶性状(E-G)与土壤环境因子的冗余分析。BD: 土壤容重; SOC: 土壤有机碳含量; TN: 土壤全氮含量; TP: 土壤全磷含量; SC : SN: 土壤碳氮比; SN : SP: 土壤氮磷比; SC : SP: 土壤碳磷比; SWC: 土壤含水量; ST: 土壤温度; LA: 叶面积; SLA: 比叶面积; LTD: 叶组织密度; LDMC: 叶干物质含量; LCC: 叶片碳含量; LNC: 叶片氮含量; LPC: 叶片磷含量; LC : LN: 叶片碳氮比; LN : LP: 叶片氮磷比; LC : LP: 叶片碳磷比; D1: 半固定沙地; D2: 固定沙地; D3: 土壤结皮固定沙地; D4: 草本植物固定沙地。箭头代表土壤环境因子, 圆圈代表植物叶性状。

Fig. 2 Redundancy analysis biplots showing the distribution of leaf trait assemblages in relation to soil environmental factors (A-C), and the relationships between individual leaf traits and soil environmental factors (E-G), across different plant functional groups and sand-fixation stages. BD, Soil bulk density; SOC, Soil organic carbon content; TN, Soil total nitrogen content; TP, Soil total phosphorus content; SC : SN, Soil carbon to nitrogen ratio; SN : SP, Soil nitrogen to phosphorus ratio; SC : SP, Soil carbon to phosphorus ratio; SWC, Soil water content; ST, Soil temperature; LA, Leaf area; SLA, Specific leaf area; LTD, Leaf tissue density; LDMC, Leaf dry matter content; LCC, Leaf carbon content; LNC, Leaf nitrogen content; LPC, Leaf phosphorus content; LC : LN, Leaf carbon to nitrogen ratio; LN : LP, Leaf nitrogen to phosphorus ratio; LC : LP, Leaf carbon to phosphorus ratio; D1, Semi-fixed dune; D2, Fixed dune; D3, Fixed dune with biological soil crusts; D4, Fixed dune with abundant herbaceous plants. Arrows indicate soil environmental factors; circles denote plant leaf traits.

表4 基于冗余分析的土壤环境因子对3种植物功能群叶性状变化的解释率

Table 4 Variance in leaf traits across three plant functional groups explained by soil environmental factors: a redundancy analysis (RDA)

土壤环境因子 Soil environmental factors	禾本科 Poaceae		非禾本科 Non-Poaceae		灌木 Shrub
土壤环境因子 Soil environmental factors	R²	P	R²	P	R²	P
土壤碳氮比 Soil carbon to nitrogen ratio (SC : SN)	0.54	< 0.001	0.30	< 0.001	0.47	< 0.001
土壤容重 Soil bulk density (BD)	0.31	< 0.001	0.26	< 0.001	0.21	< 0.001
土壤有机碳含量 Soil organic carbon content (SOC)	0.30	< 0.001	0.19	< 0.001	0.34	< 0.001
土壤全氮含量 Soil total nitrogen content (TN)	0.19	< 0.001	0.45	< 0.001	0.27	< 0.001
土壤全磷含量 Soil total phosphorus content (TP)	0.20	< 0.001	0.08	< 0.005	0.18	< 0.001
土壤含水量 Soil water content (SWC)	0.20	< 0.001	0.17	< 0.001	0.21	< 0.001
土壤氮磷比 Soil nitrogen to phosphorus ratio (SN : SP)	0.19	< 0.001	0.12	< 0.001	0.27	< 0.001
土壤温度 Soil temperature (ST)	0.07	0.054	0.12	< 0.001	0.03	0.264
土壤碳磷比 Soil carbon to phosphorus ratio (SC : SP)	0.02	0.389	0.00	0.907	0.00	0.911

图3 土壤环境因子对植物叶性状影响的相对重要性。方差分解和层次分割分析结果使用UpSet图来呈现。(A)展示了各组分(环境因子单独效应及共同效应)解释的变差百分比。(B)右侧点阵图中, 每行对应一个环境因子; 每列的孤立黑点表示各环境因子的边际效应, 多点间连线表示这些环境因子间的共同效应。左侧柱形图为各环境因子的单独效应(来自层次分割), 其值等同于该环境因子的边际效应加上与其他环境因子的共同效应的平均分配值。BD: 土壤容重; SOC: 土壤有机碳含量; TN: 土壤全氮含量; TP: 土壤全磷含量; SC : SN: 土壤碳氮比; SN : SP: 土壤氮磷比; SC : SP: 土壤碳磷比; SWC: 土壤含水量; ST: 土壤温度。***, P < 0.001。

Fig. 3 UpSet matrix layout of variation partitioning and hierarchical partitioning analysis results to show the relative importance of soil environmental factors on variation of plant traits. (A) Percentage of variation explained by each component (individual effects and common effects of environmental factors). (B) In the dot-matrix plot on the right, each row corresponds to an environmental factor. For each column, the isolated black dot represents the marginal effect of each environmental factor, lines connecting multiple dots represent the common effect among these corresponding environmental factors. Column diagram on the left shows individual effect of each environmental factor (from hierarchical partitioning), its value is equal to its marginal effect plus its average shared common effect with other environmental factors. BD, Soil bulk density; SOC, Soil organic carbon content; TN, Soil total nitrogen content; TP, Soil total phosphorus content; SC : SN, Soil carbon to nitrogen ratio; SN : SP, Soil nitrogen to phosphorus ratio; SC : SP, Soil carbon to phosphorus ratio; SWC, Soil water content; ST, Soil temperature. ***, P < 0.001.

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土壤环境因子对荒漠植物叶性状的驱动机制

How soil environmental factors shape leaf traits in arid-land vegetation

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