高山灌木扩张梯度下植物叶片与根系酚类物质的响应特征

doi:10.17521/cjpe.2025.0182

植物生态学报 ›› 2025, Vol. 49 ›› Issue (12): 2119-2136.DOI: 10.17521/cjpe.2025.0182 cstr: 32100.14.cjpe.2025.0182

高山灌木扩张梯度下植物叶片与根系酚类物质的响应特征

宋思宇¹^,², 杜飘¹^,², 林琴², 齐祥², 杜柯芋¹^,², 李聪¹^,², 陈亚梅⁵, 黄尤优⁵, 刘洋¹^,²^,³^,⁴^,^*()()

¹长江上游森林生态与保育四川省重点实验室, 成都 611130
²四川农业大学林学院, 成都 611130
³四川峨眉山森林生态系统国家定位观测研究站, 四川乐山 614000
⁴高山生态系统长期定位研究站, 四川阿坝 624000
⁵西南野生动植物资源保护教育部重点实验室, 四川南充 637009

收稿日期:2025-05-20 接受日期:2025-08-25 出版日期:2025-12-20 发布日期:2025-12-26
通讯作者: *刘洋(sicauliuyang@163.com)，ORCID：0000-0002-4795-9169
基金资助:
国家自然科学基金(32371850);国家自然科学基金(41901224);四川省科技计划资助项目(2025YFHZ0168)

Response characteristics of phenolic compounds in plant leaves and roots along an alpine shrub encroachment gradient

SONG Si-Yu¹^,², DU Piao¹^,², LIN Qin², QI Xiang², DU Ke-Yu¹^,², LI Cong¹^,², CHEN Ya-Mei⁵, HUANG You-You⁵, LIU Yang¹^,²^,³^,⁴^,^*()()

¹Forest Ecology and Conservation in the Upper Reaches of the Yangtze River Key Laboratory of Sichuan Province, Chengdu 611130, China
²College of Forestry, Sichuan Agricultural University, Chengdu 611130, China
³Sichuan Mt. Emei Forest Ecosystem National Observation and Research Station, Leshan, Sichuan 614000, China
⁴Long-term Research Station of Alpine Ecosystems, Aba, Sichuan 624000, China
⁵Key Laboratory of Southwest China Wildlife Resources Conservation, Ministry of Education, Nanchong, Sichuan 637009, China

Received:2025-05-20 Accepted:2025-08-25 Online:2025-12-20 Published:2025-12-26
Supported by:
National Natural Science Foundation of China(32371850);National Natural Science Foundation of China(41901224);Sichuan Science and Technology Program(2025YFHZ0168)

摘要/Abstract

摘要：

随着气候变化的加剧, 全球高海拔和高纬度地区的灌木扩张现象日益显著。然而, 高山地区灌木扩张过程中植物防御性化学物质(如总酚和缩合单宁)含量的变化规律及其驱动机制尚不明确。该研究以青藏高原东缘的川西高山灌丛-草甸交错带为研究区域, 沿灌木扩张梯度选取对照(盖度0%)、轻度扩张(盖度<30%)、轻中度扩张(盖度30%-45%)、中重度扩张(盖度45%-60%)和重度扩张(盖度>60%)样地, 采用空间代替时间的方法, 分析了灌木扩张对植物防御性化学物质含量的影响及其与环境因子(如海拔、土壤pH、含水量等)的关系。结果表明: (1)灌木扩张显著增加了川西高山灌丛-草甸交错带的灌丛数量、高度、体积和加权密度; (2)与草本植物和红毛花楸(Sorbus rufopilosa)相比, 高山杜鹃(Rhododendron lapponicum)在灌木扩张过程中表现出较高的酚类物质含量; (3)群落水平物种组成的变化对植物酚类物质变异的影响显著高于物种内个体间表型可塑性的种内变异; (4)植物叶片与根系的酚类物质含量均与其碳氮比显著相关, 而土壤因子影响较小, 支持了碳-养分平衡假说。该研究揭示了灌木扩张对植物防御物质的正向影响, 并为全球变化背景下的生态系统管理和植物保护提供了重要的理论依据。

关键词: 灌木扩张, 酚类物质, 种内性状变异, 碳氮比, 碳-养分平衡假说

Abstract:

Aims With the intensification of climate change, shrub expansion or encroachment in high-altitudes and high-latitude has become increasingly significant. However, the patterns and driving mechanisms of changes in plant defensive chemicals such as total phenols and condensed tannins during such shrub expansion processes in alpine regions remain poorly understood.

Methods Taking the alpine shrub-meadow transition zone or ecotone in western Sichuan on the eastern Qingzang Plateau as the study area, sample plots were established along a shrub expansion gradient including control (0% coverage), light expansion (<30% coverage), light-moderate expansion (30%-45% coverage), moderate-heavy expansion (45%-60% coverage), and heavy expansion (>60% coverage). The impact of shrub expansion on the content of plant defensive chemical compounds and their relationships with environmental factors such as altitude, soil pH, and moisture content was analyzed by a space-for-time substitution approach.

Important findings The results showed that: (1) Shrub expansion significantly increased the density, height, volume, and weighted density of shrubs in the alpine shrub-meadow transition zone of western Sichuan; (2) Compared to herbaceous plants and Sorbus rufopilosa, Rhododendron lapponicum exhibited higher levels of phenolic compounds during the shrub expansion process; (3) Changes in species composition at the community level had a significantly greater impact on the variation in plant phenolic compounds than intraspecific variation resulting from phenotypic plasticity among individuals; (4) Concentrations of phenolic compounds in both plant leaves and roots showed significant correlations with their carbon to nitrogen ratios, while soil factors had a relatively minimal effect, supporting the Carbon-nutrient Balance Hypothesis. This study sheds light on the positive impact of shrub expansion on plant defense compounds and provides critically theoretical underpinnings for ecosystem management and plant conservation in the context of global change.

Key words: shrub encroachment, phenolic compounds, intraspecific trait variation, carbon to nitrogen ratio, Carbon-nutrient Balance Hypothesis

宋思宇, 杜飘, 林琴, 齐祥, 杜柯芋, 李聪, 陈亚梅, 黄尤优, 刘洋. 高山灌木扩张梯度下植物叶片与根系酚类物质的响应特征. 植物生态学报, 2025, 49(12): 2119-2136. DOI: 10.17521/cjpe.2025.0182

SONG Si-Yu, DU Piao, LIN Qin, QI Xiang, DU Ke-Yu, LI Cong, CHEN Ya-Mei, HUANG You-You, LIU Yang. Response characteristics of phenolic compounds in plant leaves and roots along an alpine shrub encroachment gradient. Chinese Journal of Plant Ecology, 2025, 49(12): 2119-2136. DOI: 10.17521/cjpe.2025.0182

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2025.0182

https://www.plant-ecology.com/CN/Y2025/V49/I12/2119

图/表 9

表1 高山灌丛-草甸分布区不同灌木扩张水平下样方基本特征

Table 1 Basic characteristics of sample plots under different shrub expansion levels in alpine shrub-meadow distribution areas

灌木扩张水平 Shrub encroachment level	冠幅 Crown width (m²)	盖度 Coverage (%)	高度 Height (m)	灌丛数量 Number of shrubs	灌丛体积 Shrub volume (m³)	灌丛加权密度 Weighted shrub density	海拔 Altitude (m)
对照(高山草甸) CK (alpine meadow)	0.00 (0.00)^e	0.00 (0.00)^e	0.00 (0.00)^c	0.00 (0.00)^c	0.00 (0.00)^e	0.00 (0.00)^c	4 200.00 (0.00)^a
轻度扩张 SE-I	106.31 (6.85)^d	26.58 (1.71)^d	1.44 (0.15)^b	49.80 (6.59)^b	77.14 (10.29)^d	653.33 (79.74)^b	4 110.00 (18.71)^b
轻中度扩张 SE-II	153.35 (7.45)^c	38.34 (1.86)^c	1.88 (0.19)^ab	59.80 (5.64)^b	130.51 (12.35)^c	815.21 (67.40)^b	4 090.00 (18.71)^b
中重度扩张 SE-III	212.50 (10.93)^b	53.12 (2.73)^b	1.68 (0.13)^ab	72.00 (12.02)^ab	175.71 (16.99)^b	978.73 (158.74)^ab	4 080.00 (33.91)^b
重度扩张 SE-IV	280.47 (17.70)^a	70.12 (4.43)^a	1.97 (0.19)^a	92.40 (14.92)^a	281.87 (20.22)^a	1 288.15 (197.77)^a	4 020.00 (12.25)^c
F	105.88	105.87	29.18	13.46	58.58	15.26	10.63
p	<0.001	<0.001	<0.001	<0.001	<0.001	<0.001	<0.001

表2 不同灌木扩张水平下高山灌丛-草甸分布区植物盖度特征(平均值±标准误, n = 5)

Table 2 Plant coverage characteristics under different shrub expansion levels in alpine shrub-meadow distribution areas (mean ± SE, n = 5)

灌木 Shrub	对照 CK	轻度扩张 SE-I	轻中度扩张 SE-II	中重度扩张 SE-III	重度扩张 SE-IV
高山杜鹃 Rhododendron lapponicum	-	11.16 ± 2.53^Aa	11.66 ± 4.04^Aa	21.04 ± 7.86^Aa	6.00 ± 1.88^BCa
红毛花楸 Sorbus rufopilosa	-	7.70 ± 1.78^Ab	13.01 ± 3.83^Ab	22.30 ± 8.21^Ab	41.19 ± 3.41^Aa
金露梅 Dasiphora fruticosa	-	2.95 ± 0.62^Bab	3.76 ± 1.41^Bab	0.79 ± 0.37^Bb	5.73 ± 1.55^BCa
软条七蔷薇 Rosa henryi	-	2.32 ± 0.84^Bb	4.45 ± 1.22^Bb	2.55 ± 0.94^Bb	8.51 ± 2.06^Ba
绣线菊 Spiraea salicifolia	-	0.23 ± 0.21^Ba	0.18 ± 0.18^Ba	0.40 ± 0.37^Ba	1.37 ± 1.00^Ca
高山柳 Salix takasagoalpina	-	0.97 ± 0.36^Ba	2.65 ± 1.48^Ba	3.52 ± 1.68^Ba	2.94 ± 1.30^BCa
华西小檗 Berberis silva-taroucana	-	0.03 ± 0.03^Ba	0.51 ± 0.19^Ba	0.58 ± 0.28^Ba	0.11 ± 0.08^Ca
F	-	11.56	5.05	5.13	58.56
p	-	<0.001	<0.01	<0.01	<0.001

表3 不同灌木扩张水平下植物酚类物质及碳氮含量的三因素/双因素方差分析(F值)

Table 3 Three-way/two-way ANOVA of phenolic compounds and carbon-nitrogen content under different shrub expansion levels (F value)

变异来源 Source of variation	总酚 Total phenolic	缩合单宁 Condensed tannin	碳 Carbon	氮 Nitrogen	碳氮比 Carbon:nitrogen	总酚RII Total phenolic RII	缩合单宁RII Condensed tannin RII
灌木扩张水平 SE	1.98^ns	2.18^ns	0.94^ns	1.24^ns	1.16^ns	0.30^ns	2.03^ns
植物类群 Plant group	136.32^***	117.79^***	2.99^ns	29.10^***	72.88^***	1.66^ns	2.87^ns
器官类型 Organ	404.94^***	112.01^***	69.67^***	127.36^***	83.50^***	-	-
灌木扩张水平×植物类群 SE × Plant group	1.17^ns	2.25^*	1.10^ns	1.02^ns	1.03^ns	-	-
灌木扩张水平×器官类型 SE × Organ	1.39^ns	0.84^ns	0.33^ns	1.66^ns	0.22^ns	0.42^ns	0.14^ns
植物类群×器官类型 Plant group × Organ	76.90^***	63.33^***	0.21^ns	9.36^***	4.04^*	-	-
灌木扩张水平×植物类群×器官类型 SE × Plant group × Organ	0.72^ns	1.73^ns	0.77^ns	1.24^ns	1.29^ns	-	-

表4 不同灌木扩张水平下植物不同器官(叶片、根系)碳氮含量及其比值

Table 4 Carbon and nitrogen content and their ratios in different plant organs (leaves, roots) under different shrub expansion levels

灌木扩张水平 Shrub encroachment level	植物类群 Plant group	叶片碳 Leaf C (%)	叶片氮 Leaf N (%)	叶片碳氮比 Leaf C:N	根系碳 Root C (%)	根系氮 Root N (%)	根系碳氮比 Root C:N
对照(高山草甸) CK (alpine meadow)	草本植物 Herb	29.27 (2.23)^a	2.72 (0.21)^a	10.78 (0.07)^b	19.91 (1.36)^a	0.87 (0.06)^a	23.03 (1.62)^a
轻度扩张 SE-I	草本植物 Herb	31.95 (2.70)^Aa	1.96 (0.22)^Aa	16.68 (1.25)^Ba	25.41 (4.97)^Aa	0.88 (0.15)^Aa	29.68 (4.37)^Ba
	高山杜鹃 Rhododendron lapponicum	35.29 (0.93)^Aa	0.88 (0.07)^Bb	41.24 (4.26)^Aa	25.81 (2.04)^Aa	0.60 (0.04)^Aab	43.08 (2.59)^Aa
	红毛花楸 Sorbus rufopilosa	31.55 (0.41)^Aa	2.18 (0.14)^Aa	14.67 (0.69)^Ba	22.76 (2.69)^Aa	0.66 (0.02)^Aa	34.43 (3.50)^ABa
轻中度扩张 SE-II	草本植物 Herb	38.02 (4.99)^Aa	2.73 (0.49)^Aa	14.71 (1.48)^Ba	23.38 (1.52)^Aa	0.99 (0.27)^Aa	28.26 (5.34)^Ba
	高山杜鹃 R. lapponicum	36.76 (1.79)^Aa	1.16 (0.10)^Ba	32.43 (2.86)^Aa	25.31 (2.10)^Aa	0.54 (0.04)^Ab	48.09 (4.44)^Aa
	红毛花楸 S. rufopilosa	28.98 (2.18)^Aa	2.12 (0.30)^ABa	14.46 (1.47)^Ba	24.40 (1.87)^Aa	0.87 (0.05)^Aa	28.51 (2.80)^Ba
中重度扩张 SE-III	草本植物 Herb	30.12 (1.72)^Aa	2.18 (0.05)^Aa	13.81 (0.64)^Bab	21.45 (2.07)^Aa	1.64 (0.81)^Aa	20.85 (4.46)^Ba
	高山杜鹃 R. lapponicum	33.36 (1.33)^Aa	0.90 (0.01)^Bb	36.96 (1.69)^Aa	26.31 (2.09)^Aa	0.63 (0.04)^Aab	42.41 (3.92)^Aa
	红毛花楸 S. rufopilosa	33.81 (2.31)^Aa	2.30 (0.18)^Aa	14.78 (0.27)^Ba	23.91 (4.79)^Aa	0.67 (0.11)^Aa	36.94 (5.56)^Aa
重度扩张 SE-IV	草本植物 Herb	28.60 (1.58)^Aa	1.93 (0.17)^Aa	15.24 (1.38)^Ba	21.12 (2.46)^Aa	0.76 (0.06)^Aa	27.98 (3.04)^Aa
	高山杜鹃 R. lapponicum	32.68 (1.51)^Aa	1.04 (0.08)^Bab	32.01 (2.09)^Aa	26.96 (2.30)^Aa	0.66 (0.02)^Aa	40.67 (2.60)^Aa
	红毛花楸 S. rufopilosa	32.33 (3.13)^Aa	1.92 (0.16)^Aa	17.83 (3.52)^Ba	24.84 (1.37)^Aa	0.98 (0.19)^Aa	30.89 (7.25)^Aa

表4 不同灌木扩张水平下植物不同器官(叶片、根系)碳氮含量及其比值

Table 4 Carbon and nitrogen content and their ratios in different plant organs (leaves, roots) under different shrub expansion levels

灌木扩张水平 Shrub encroachment level	植物类群 Plant group	叶片碳 Leaf C (%)	叶片氮 Leaf N (%)	叶片碳氮比 Leaf C:N	根系碳 Root C (%)	根系氮 Root N (%)	根系碳氮比 Root C:N
对照(高山草甸) CK (alpine meadow)	草本植物 Herb	29.27 (2.23)^a	2.72 (0.21)^a	10.78 (0.07)^b	19.91 (1.36)^a	0.87 (0.06)^a	23.03 (1.62)^a
轻度扩张 SE-I	草本植物 Herb	31.95 (2.70)^Aa	1.96 (0.22)^Aa	16.68 (1.25)^Ba	25.41 (4.97)^Aa	0.88 (0.15)^Aa	29.68 (4.37)^Ba
	高山杜鹃 Rhododendron lapponicum	35.29 (0.93)^Aa	0.88 (0.07)^Bb	41.24 (4.26)^Aa	25.81 (2.04)^Aa	0.60 (0.04)^Aab	43.08 (2.59)^Aa
	红毛花楸 Sorbus rufopilosa	31.55 (0.41)^Aa	2.18 (0.14)^Aa	14.67 (0.69)^Ba	22.76 (2.69)^Aa	0.66 (0.02)^Aa	34.43 (3.50)^ABa
轻中度扩张 SE-II	草本植物 Herb	38.02 (4.99)^Aa	2.73 (0.49)^Aa	14.71 (1.48)^Ba	23.38 (1.52)^Aa	0.99 (0.27)^Aa	28.26 (5.34)^Ba
	高山杜鹃 R. lapponicum	36.76 (1.79)^Aa	1.16 (0.10)^Ba	32.43 (2.86)^Aa	25.31 (2.10)^Aa	0.54 (0.04)^Ab	48.09 (4.44)^Aa
	红毛花楸 S. rufopilosa	28.98 (2.18)^Aa	2.12 (0.30)^ABa	14.46 (1.47)^Ba	24.40 (1.87)^Aa	0.87 (0.05)^Aa	28.51 (2.80)^Ba
中重度扩张 SE-III	草本植物 Herb	30.12 (1.72)^Aa	2.18 (0.05)^Aa	13.81 (0.64)^Bab	21.45 (2.07)^Aa	1.64 (0.81)^Aa	20.85 (4.46)^Ba
	高山杜鹃 R. lapponicum	33.36 (1.33)^Aa	0.90 (0.01)^Bb	36.96 (1.69)^Aa	26.31 (2.09)^Aa	0.63 (0.04)^Aab	42.41 (3.92)^Aa
	红毛花楸 S. rufopilosa	33.81 (2.31)^Aa	2.30 (0.18)^Aa	14.78 (0.27)^Ba	23.91 (4.79)^Aa	0.67 (0.11)^Aa	36.94 (5.56)^Aa
重度扩张 SE-IV	草本植物 Herb	28.60 (1.58)^Aa	1.93 (0.17)^Aa	15.24 (1.38)^Ba	21.12 (2.46)^Aa	0.76 (0.06)^Aa	27.98 (3.04)^Aa
	高山杜鹃 R. lapponicum	32.68 (1.51)^Aa	1.04 (0.08)^Bab	32.01 (2.09)^Aa	26.96 (2.30)^Aa	0.66 (0.02)^Aa	40.67 (2.60)^Aa
	红毛花楸 S. rufopilosa	32.33 (3.13)^Aa	1.92 (0.16)^Aa	17.83 (3.52)^Ba	24.84 (1.37)^Aa	0.98 (0.19)^Aa	30.89 (7.25)^Aa

图1 不同灌木扩张水平下植物不同器官(叶片、根系)总酚含量。对照, 高山草甸; Herb, 草本植物; Rhododendron lapponicum, 高山杜鹃; Sorbus rufopilosa, 红毛花楸。不同小写字母表示同一个植物类群在不同灌木扩张水平下差异显著(单因素方差分析, p < 0.05), 不同大写字母表示同一灌木扩张水平下不同植物类群差异显著(单因素方差分析, p < 0.05)。

Fig. 1 Variations in total phenolics content in different plant organs (leaves, roots) under different shrub expansion levels. CK, alpine meadow. Different lowercase letters indicate significant differences in the same plant group across different shrub expansion levels (one-way ANOVA, p < 0.05), while different uppercase letters represent significant differences among plant groups within the same shrub expansion level (one-way ANOVA, p < 0.05). SE-I, light shrub expansion; SE-II, light-moderate shrub expansion; SE-III, moderate-heavy shrub expansion; SE-IV, heavy shrub expansion.

图2 不同灌木扩张水平下植物不同器官(叶片、根系)缩合单宁含量。对照, 高山草甸; Herb, 草本植物; Rhododendron lapponicum, 高山杜鹃; Sorbus rufopilosa, 红毛花楸。不同小写字母表示同一个植物类群在不同灌木扩张水平下差异显著(单因素方差分析, p < 0.05), 不同大写字母表示同一灌木扩张水平下不同植物类群差异显著(单因素方差分析, p < 0.05)。

Fig. 2 Variations in condensed tannins (CT) content in different plant organs (leaves, roots) under different shrub expansion levels. CK, alpine meadow. Different lowercase letters indicate significant differences in the same plant group across different shrub expansion levels (one-way ANOVA, p < 0.05), while different uppercase letters represent significant differences among plant groups within the same shrub expansion level (one-way ANOVA, p < 0.05). SE-I, light shrub expansion; SE-II, light-moderate shrub expansion; SE-III, moderate-heavy shrub expansion; SE-IV, heavy shrub expansion.

图3 不同灌木扩张水平下植物不同器官(叶片、根系)酚类物质相对作用指数(RII)。*, 灌木扩张组与对照组(未扩张高山草甸)独立t检验差异显著(ns, p > 0.05; *, p < 0.05; **, p < 0.01; ***, p < 0.001)。CT, 缩合单宁。

Fig. 3 Variations in relative interaction index (RII) of phenolic compounds in different plant organs (leaves, roots) under different shrub encroachment levels. *, indicates a significant difference between the shrub encroachment group and the control group (non-encroached alpine meadow) (ns, not significant; *, p < 0.05; **, p < 0.01; ***, p < 0.001). CT, condensed tannins. SE-I, light shrub expansion; SE-II, light-moderate shrub expansion; SE-III, moderate-heavy shrub expansion; SE-IV, heavy shrub expansion.

图4 灌木扩张水平下根系总酚、叶片总酚、根系缩合单宁和叶片缩合单宁含量的固定平均值、特定平均值及种内性状变异效应。固定平均值基于跨处理条件下不变的物种性状值, 特定平均值基于不同扩张水平下各性状的特定值。误差条表示标准误差。CK, 高山草甸; SE-I, 轻度灌木扩张; SE-II, 轻中度灌木扩张; SE-III, 中重度灌木扩张; SE-IV, 重度灌木扩张。CT, 缩合单宁。采用单因素方差分析检验不同灌木扩张水平间固定平均值、特定平均值及种内性状变异效应的差异。

Fig. 4 Variations in fixed averages (FA) and specific averages (SA), and intraspecific variability effects (IVE) on the content of total root phenol, total leaf phenol, root condensed tannin, and leaf condensed tannin across different shrub expansion levels. Fixed averages are based on species trait values that remain constant across treatments, while specific averages are based on the trait values specific to each expansion level. CK, alpine meadow; SE-I, light shrub expansion; SE-II, light-moderate shrub expansion; SE-III, moderate-heavy shrub expansion; SE-IV, heavy shrub expansion. CT, condensed tannins. The differences in the effects of fixed averages, specific averages, and intraspecific variability effects across different shrub encroachment levels were tested using One-way ANOVA.

图5 环境因子对植物不同器官(叶片、根系)酚类物质含量的影响——基于偏最小二乘(PLS)分析(平均值±标准误)。PLS系数大于0表示正向影响, 系数小于0表示负向影响。R2代表模型拟合度。Alt, 海拔; CT, 缩合单宁; Leaf_C, 叶片碳含量; Leaf_N, 叶片氮含量; Leaf_C:N, 叶片碳氮比; Root_C, 根系碳含量; Root_N, 根系氮含量; Root_C:N, 根系碳氮比; Shrub_C, 灌丛盖度; Shrub_H, 灌丛高度; Shrub_L, 灌木扩张水平; Shrub_N, 灌丛数量; Shrub_V, 灌丛体积; Shrub_W, 灌丛加权密度; Soil_AP, 土壤速效磷含量; Soil_C, 土壤有机碳含量; Soil_C:N, 土壤碳氮比; Soil_MC, 土壤含水量; Soil_N, 土壤全氮含量; Soil_pH, 土壤pH。

Fig. 5 Effects of environmental factors on phenolic compound contents in different plant organs (leaves, roots) were quantified using partial least squares (PLS) analysis (mean ± SE). PLS coefficients > 0 indicate positive effects, while those < 0 indicate negative effects. R2 values represent the goodness of model fit. Alt, altitude; CT, condensed tannins; Leaf_C, leaf carbon content; Leaf_N, leaf nitrogen content; Leaf_C:N, leaf carbon-to-nitrogen ratio; Root_C, root carbon content; Root_N, root nitrogen content; Root_C:N, root carbon-to-nitrogen ratio; Shrub_C, shrub coverage; Shrub_H, shrub height; Shrub_L, shrub encroachment level; Shrub_N, number of shrubs; Shrub_V, shrub volume; Shrub_W, weighted shrub density; Soil_AP, soil available phosphorus content; Soil_C, soil organic carbon content; Soil_C:N, soil carbon-to-nitrogen ratio; Soil_MC, soil moisture content; Soil_N, soil total nitrogen content; Soil_pH, soil pH. VIP, value of importance.

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高山灌木扩张梯度下植物叶片与根系酚类物质的响应特征

Response characteristics of phenolic compounds in plant leaves and roots along an alpine shrub encroachment gradient

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