三峡回水扰动增强入库小流域河岸带典型草本植物功能性状的局域分化

doi:10.17521/cjpe.2024.0394

植物生态学报 ›› 2026, Vol. 50 ›› Issue (2): 442-460.DOI: 10.17521/cjpe.2024.0394 cstr: 32100.14.cjpe.2024.0394

三峡回水扰动增强入库小流域河岸带典型草本植物功能性状的局域分化

戴丽君¹^,²^,³, 向玲艺⁴, 蹇陈¹^,²^,³, 王晓锋¹^,²^,³^,^*()

¹ 重庆师范大学地理与旅游学院, 重庆 401331
² 山区生态系统碳循环与碳调控重庆市重点实验室, 重庆 401331
³ 三峡库区地表生态过程重庆市野外科学观测研究站, 重庆 405400
⁴ 重庆地质矿产研究院, 重庆 401120

收稿日期:2024-11-05 接受日期:2025-05-01 出版日期:2026-02-28 发布日期:2026-04-01
通讯作者: *王晓锋(xiaofeng6540@163.com)
基金资助:
国家自然科学基金(32371680);重庆市教委科学技术研究项目(KJQN20220053)

Intensified local differentiation in plant functional traits among typical herbaceous plants by backwater disturbance within riparian zone of small watersheds in the Three Gorges Reservoir area

DAI Li-Jun¹^,²^,³, XIANG Ling-Yi⁴, JIAN Chen¹^,²^,³, WANG Xiao-Feng¹^,²^,³^,^*()

¹ College of Geography and Tourism, Chongqing Normal University, Chongqing 401331, China
² Chongqing Key Laboratory of Carbon Cycle and Carbon Regulation in Mountainous Ecosystems, Chongqing 401331, China
³ Three Gorges Reservoir Area Earth Surface Ecological Processes of Chongqing Observation and Research Station, Chongqing 405400, China
⁴ Chongqing Institute of Geology and Mineral Resources, Chongqing 401120, China

Received:2024-11-05 Accepted:2025-05-01 Online:2026-02-28 Published:2026-04-01
Contact: *WANG Xiao-Feng (xiaofeng6540@163.com)
Supported by:
National Natural Science Foundation of China(32371680);Science and Technology Research Project of Chongqing Education Commission(KJQN20220053)

摘要/Abstract

摘要：

三峡水库蓄水导致大量自然河岸带转变为反季节水位涨落的水库消落带, 形成了局域范围内生境的彻底转变, 对植物性状分化产生重要影响。该研究旨在探讨三峡回水扰动下入库小流域“河岸-消落带”生境转变对适生草本植物功能性状分异的影响及植物的适应策略。该研究以三峡库区入库小流域——白家溪为例, 选取了狗牙根(Cynodon dactylon)、狼杷草(Bidens tripartita)、苍耳(Xanthium strumarium)、水蓼(Polygonum hydropiper)等4种主要适生草本植物为对象, 分析了上游河岸带向下游消落带生境转变中17个植物功能性状指标的空间分异特征, 采用Pearson相关性分析植物各功能性状间的协同演变关联, 利用冗余分析分析生境因子对植物功能性状分异的影响。主要结果: (1)随着自然河岸带向消落带转变, 4种植物形态性状变异明显, 植株高度均增加, 根长变短, 这主要与消落带生境的生长期限制以及植物群落结构相对单一、竞争减弱有关; 同时, 大部分植物叶片厚度增加, 叶面积增大, 主要受夏季消落带干旱环境加剧的驱动; 三峡回水扰动显著增加了物种在局域范围内的形态变异。(2) 4种植物叶片的叶绿素含量和净光合速率在消落带明显高于自然河岸带, 表明植物在消落带生境的生长期限制下可能形成了光合效率提高、生长速度加快的适应策略。(3)在三峡回水的环境筛选作用下, 4种植物形成了独特的“生长-耐旱”的权衡策略, 并表现出明显的趋同适应; 但狗牙根的性状变异性较强, 尤其是其光合性状变异性强于其他3种植物, 表明狗牙根对生境转变具有更强的适应能力; 相比之下, 狼杷草、苍耳和水蓼形成了较为保守的适应对策。(4)冗余分析显示, 4种植物功能性状的变异特征在上游自然河岸带主要与土壤水分含量有关; 而在中游过渡带和下游消落带, 则主要与淹水深度、土壤pH、土壤温度有关, 表明淹水环境差异对植物功能性状的分异起到了促进作用。

关键词: 草本植物, 功能性状, 适应策略, 消落带, 三峡库区

Abstract:

Aims The impoundment of the Three Gorges Reservoir (TGR) has led to the transformation of a large number of natural riparian zones into reservoir drawdown zones, characterized by reverse seasonal water level fluctuations. This transformation has fundamentally altered the local habitat, significantly impacting the differentiation of plant morphological traits. This study aims to reveal the impact of habitat transformation from “riverbank to drawdown zone” as a result of the TGR backwater disturbance on the functional trait differentiation of adapted herbaceous plants as well as their adaptation strategies.

Methods This study used a typical small watershed (Baijiaxi) in TGR area as a case. Four herbaceous plant species (Cynodon dactylon, Bidens tripartita, Xanthium strumarium, and Polygonum hydropiper) suitable for the local environment were selected as objects. The spatial differentiation characteristics of 17 plant functional trait indicators were analyzed across the habitat transformation from the riparian zone to the drawdown zone. Pearson correlation analysis was employed to explore the co-evolutionary relationships among functional traits, while redundancy analysis (RDA) was conducted to assess the influence of habitat factors on the differentiation of these traits.

Important findings (1) The transition from natural riparian habitats into the drawdown zone resulted in significant variations in the morphological traits of the four plant species. These variations were characterized by increased plant height and decreased root length. These changes were primarily attributed to the restricted growth period imposed by water-level fluctuations and the relatively simplified plant community structure with reduced competitive pressure. Concurrently, leaf thickness increased, and leaf area expanded in most plants, which was mainly driven by intensified drought stress during summer in the drawdown zone. The results highlighted that the backwater effects of the TGR significantly amplified local-scale shape variations among species. (2) The chlorophyll content and net photosynthetic rate of leaves in the four plant species were significantly higher in the drawdown zone compared to those in the natural riparian zone, indicating that plants may have developed an adaptation strategy to enhance photosynthetic efficiency and accelerate growth rates in response to habitat limitations. (3) Under the habitat screening effect of the backwater in the TGR, the four plant species developed a trade-off strategy between “growth” and “drought tolerance”, showing convergent adaptation strategies. Cynodon dactylon exhibited stronger phenotypic variability, especially in photosynthetic traits, suggesting greater adaptability to habitat changes. In contrast, Bidens tripartita, Xanthium strumarium, and Polygonum hydropiper formed more conservative adaptation strategies. (4) RDA showed that variations in functional traits of the four plant species within the upstream riparian zone were primarily associated with soil moisture. While, in the midstream and downstream drawdown zone, these variations were predominantly linked to flooding depth, soil pH and soil temperature, indicating that differences in flooding regimes drove the differentiation of plant functional traits.

Key words: herbaceous plant species, functional trait, adaptive strategy, drawdown zone, Three Gorges Reservoir

戴丽君, 向玲艺, 蹇陈, 王晓锋. 三峡回水扰动增强入库小流域河岸带典型草本植物功能性状的局域分化. 植物生态学报, 2026, 50(2): 442-460. DOI: 10.17521/cjpe.2024.0394

DAI Li-Jun, XIANG Ling-Yi, JIAN Chen, WANG Xiao-Feng. Intensified local differentiation in plant functional traits among typical herbaceous plants by backwater disturbance within riparian zone of small watersheds in the Three Gorges Reservoir area. Chinese Journal of Plant Ecology, 2026, 50(2): 442-460. DOI: 10.17521/cjpe.2024.0394

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2024.0394

https://www.plant-ecology.com/CN/Y2026/V50/I2/442

图/表 10

图1 白家溪消落带采样点布设。S1, S2, 自然河岸带; S3-S5, 过渡带; S6-S8, 消落带。

Fig. 1 Layout of sampling site in the drawdown zone of Baijiaxi. S1, S2, natural riparian zone; S3-S5, transition zone; S6-S8, drawdown zone.

表1 白家溪流域自然河岸带至消落带土壤物理性状及植物群落状况

Table 1 Soil physical properties from natural riparian zone to subsidence zone in Baijiaxi Watershed

断面 Transect	土壤pH Soil pH	土壤温度 Soil temperature (℃)	土壤湿度 Soil humidity (%)	土壤含水量 Soil water content (%)	植物群落 Phytocoenosium
断面 Transect	土壤pH Soil pH	土壤温度 Soil temperature (℃)	土壤湿度 Soil humidity (%)	土壤含水量 Soil water content (%)	优势种 Dominant species	盖度 Coverage (%)
1	7.39 ± 0.37^c	34.3 ± 4.6^ab	49.7 ± 14.7^a	35.4 ± 9.8^a	水蓼、空心莲子草、狗牙根、苍耳 Polygonum hydropiper, Alternanthera philoxeroides, Cynodon dactylon, Xanthium strumarium	83-100
2	7.29 ± 0.49^c	33.0 ± 5.2^b	30.5 ± 11.6^b	20.9 ± 4.5^b	空心莲子草、狗牙根、狼杷草 Alternanthera philoxeroides, Cynodon dactylon, Bidens tripartite	88-100
3	7.84 ± 0.50^b	37.3 ± 5.4^a	22.3 ± 5.2^b	14.0 ± 3.8^c	狗牙根、苍耳、水蓼、狼杷草、香附子 Cynodon dactylon, Xanthium strumarium, Polygonum hydropiper, Bidens tripartite, Cyperus rotundus	95-100
4	7.84 ± 0.32^b	35.7 ± 3.6^ab	20.0 ± 6.0^b	12.8 ± 4.2^c	狗牙根、苍耳、水蓼、苘麻、狼杷草、香附子 Cynodon dactylon, Xanthium strumarium, Polygonum hydropiper, Abutilon theophrasti, Bidens tripartite, Cyperus rotundus	95-100
5	8.17 ± 0.53^ab	36.5 ± 3.5^ab	21.3 ± 4.3^b	10.0 ± 2.1^c	狗牙根、苍耳、苘麻、水蓼、狼杷草 Cynodon dactylon, Xanthium strumarium, Abutilon theophrasti, Polygonum hydropiper, Bidens tripartite	100
6	8.09 ± 0.26^ab	36.4 ± 2.3^ab	22.1 ± 4.9^b	7.8 ± 1.6^c	狗牙根、苍耳、苘麻、水蓼、狼杷草 Cynodon dactylon, Xanthium strumarium, Abutilon theophrasti, Polygonum hydropiper, Bidens tripartite	100
7	8.36 ± 0.4^a	38.3 ± 4.7^a	21.6 ± 3.5^b	7.9 ± 1.8^c	狗牙根、苍耳、苘麻、水蓼、狼杷草 Cynodon dactylon, Xanthium strumarium, Abutilon theophrasti, Polygonum hydropiper, Bidens tripartite	100
8	8.31 ± 0.56^a	35.6 ± 2.9^ab	21.3 ± 3.2^b	7.7 ± 0.9^c	狗牙根、苍耳、苘麻、水蓼、狼杷草 Cynodon dactylon, Xanthium strumarium, Abutilon theophrasti, Polygonum hydropiper, Bidens tripartite	100

表1 白家溪流域自然河岸带至消落带土壤物理性状及植物群落状况

Table 1 Soil physical properties from natural riparian zone to subsidence zone in Baijiaxi Watershed

断面 Transect	土壤pH Soil pH	土壤温度 Soil temperature (℃)	土壤湿度 Soil humidity (%)	土壤含水量 Soil water content (%)	植物群落 Phytocoenosium
断面 Transect	土壤pH Soil pH	土壤温度 Soil temperature (℃)	土壤湿度 Soil humidity (%)	土壤含水量 Soil water content (%)	优势种 Dominant species	盖度 Coverage (%)
1	7.39 ± 0.37^c	34.3 ± 4.6^ab	49.7 ± 14.7^a	35.4 ± 9.8^a	水蓼、空心莲子草、狗牙根、苍耳 Polygonum hydropiper, Alternanthera philoxeroides, Cynodon dactylon, Xanthium strumarium	83-100
2	7.29 ± 0.49^c	33.0 ± 5.2^b	30.5 ± 11.6^b	20.9 ± 4.5^b	空心莲子草、狗牙根、狼杷草 Alternanthera philoxeroides, Cynodon dactylon, Bidens tripartite	88-100
3	7.84 ± 0.50^b	37.3 ± 5.4^a	22.3 ± 5.2^b	14.0 ± 3.8^c	狗牙根、苍耳、水蓼、狼杷草、香附子 Cynodon dactylon, Xanthium strumarium, Polygonum hydropiper, Bidens tripartite, Cyperus rotundus	95-100
4	7.84 ± 0.32^b	35.7 ± 3.6^ab	20.0 ± 6.0^b	12.8 ± 4.2^c	狗牙根、苍耳、水蓼、苘麻、狼杷草、香附子 Cynodon dactylon, Xanthium strumarium, Polygonum hydropiper, Abutilon theophrasti, Bidens tripartite, Cyperus rotundus	95-100
5	8.17 ± 0.53^ab	36.5 ± 3.5^ab	21.3 ± 4.3^b	10.0 ± 2.1^c	狗牙根、苍耳、苘麻、水蓼、狼杷草 Cynodon dactylon, Xanthium strumarium, Abutilon theophrasti, Polygonum hydropiper, Bidens tripartite	100
6	8.09 ± 0.26^ab	36.4 ± 2.3^ab	22.1 ± 4.9^b	7.8 ± 1.6^c	狗牙根、苍耳、苘麻、水蓼、狼杷草 Cynodon dactylon, Xanthium strumarium, Abutilon theophrasti, Polygonum hydropiper, Bidens tripartite	100
7	8.36 ± 0.4^a	38.3 ± 4.7^a	21.6 ± 3.5^b	7.9 ± 1.8^c	狗牙根、苍耳、苘麻、水蓼、狼杷草 Cynodon dactylon, Xanthium strumarium, Abutilon theophrasti, Polygonum hydropiper, Bidens tripartite	100
8	8.31 ± 0.56^a	35.6 ± 2.9^ab	21.3 ± 3.2^b	7.7 ± 0.9^c	狗牙根、苍耳、苘麻、水蓼、狼杷草 Cynodon dactylon, Xanthium strumarium, Abutilon theophrasti, Polygonum hydropiper, Bidens tripartite	100

图2 白家溪4种植物叶形态沿河流纵向的空间分布特征。S1, S2, 自然河岸带; S3-S5, 过渡带; S6-S8, 消落带。不同小写字母表示不同采样点间差异显著(p < 0.05)。

Fig. 2 Spatial distribution characteristics of leaf morphology of four species in Baijiaxi along the river. S1, S2, natural riparian zone; S3-S5, transition zone; S6-S8, drawdown zone. Lowercase letters in the figure represent significant difference (p < 0.05).

图3 白家溪植物根茎形态的纵向空间分布特征(平均值±标准差)。S1, S2, 自然河岸带; S3-S5, 过渡带; S6-S8, 消落带。C为标准化后的植物根长。F为标准化后的植株高度。图中小写字母表示差异显著（p < 0.05）。

Fig. 3 Longitudinal spatial distribution characteristics of root and stem morphology of plants in Baijiaxi (mean ± SD). S1, S2, natural riparian zone; S3-S5, transition zone; S6-S8, drawdown zone. C is the plant root length after standardization. F is the plant height after standardization. Lowercase letters in the figure represent significant difference (p < 0.05).

图4 白家溪植物根茎叶鲜质量和干质量的纵向空间分布。S1, S2, 自然河岸带; S3-S5, 过渡带; S6-S8, 消落带。p < 0.05表示线性关系显著; p > 0.05表示线性关系不显著。

Fig. 4 Longitudinal spatial distribution of fresh mass and dry mass of plant roots, stems and leaves in Baijiaxi Watershed. S1, S2, natural riparian zone; S3-S5, transition zone; S6-S8, drawdown zone. p < 0.05 indicates a significant linear relationship; p > 0.05 indicates that the linear relationship is not significant.

图5 白家溪植物根茎叶相对生物量及含水量纵向空间分布。S1, S2, 自然河岸带; S3-S5, 过渡带; S6-S8, 消落带。

Fig. 5 Longitudinal zonation of relative biomass and water content in roots, stems, and leaves of Baijiaxi plants. S1, S2, natural riparian zone; S3-S5, transition zone; S6-S8, drawdown zone.

图6 白家溪植物光合性状的纵向空间分布。S1, S2, 自然河岸带; S3-S5, 过渡带; S6-S8, 消落带。p < 0.05表示线性关系显著; p > 0.05表示线性关系不显著。

Fig. 6 Longitudinal spatial distribution of photosynthetic traits in Baijiaxi plants. S1, S2, natural riparian zone; S3-S5, transition zone; S6-S8, drawdown zone. p < 0.05 indicates a significant linear relationship; p > 0.05 indicates that the linear relationship is not significant. Chl, chlorophyll; Pn, net photosynthetic rate.

表2 4种植物功能性状沿河流纵向空间分布的变异系数(%)

Table 2 Coefficient of variation (%) of 4 plants functional traits in longitudinal spatial distribution along the river

性状 Trait	狗牙根 Cynodon dactylon	苍耳 Bidens tripartita	狼杷草 Xanthium strumarium	水蓼 Polygonum hydropiper
植株高度 Plant height	25	11	7	26
根长 Root length	22	18	20	31
叶长 Leaf length	9	11	6	30
叶厚 Leaf thickness	42	24	19	48
叶面积 Leaf area	22	25	27	34
比叶面积 Specific leaf area	34	16	36	13
叶鲜质量 Leaf fresh mass	34	16	25	33
叶干质量 Leaf dry mass	43	17	20	34
茎鲜质量 Stem fresh mass	30	31	29	19
茎干质量 Stem dry mass	28	26	29	23
根鲜质量 Root fresh mass	31	34	23	16
根干质量 Root dry mass	33	41	32	14
叶绿素含量 Chlorophyll content	27	8	14	13
净光合速率 Net photosynthetic rate	29	14	22	14
叶含水量 Leaf water content	19	2	5	8
茎含水量 Stem water content	19	9	5	7
根含水量 Root water content	31	18	11	9

图7 白家溪4种植物功能性状沿河流纵向空间分布的相关性。蓝色代表正相关, 红色代表负相关, 颜色越深则表示相关关系越强。Chl, 叶绿素含量; LA, 叶面积; LDM, 叶干质量; LFM, 叶鲜质量; LL, 叶长; LT, 叶厚度; LWC, 叶含水量; PH, 植株高度; Pn, 净光合速率; RDM, 根干质量; RFM, 根鲜质量; RL, 根长; RWC, 根含水量; SDM, 茎干质量; SFM, 茎鲜质量; SLA, 比叶面积; SWC, 茎含水量。*, p < 0.05; **, p < 0.01; ***, p < 0.001。

Fig. 7 Correlation of functional traits among four plant species along the longitudinal river gradient in Baijiaxi. Blue represents a positive correlation, red represents a negative correlation, and the darker the color, the greater the correlation. Chl, chlorophyll content; LA, leaf area; LDM, leaf dry mass; LFM, leaf fresh mass; LL, leaf length; LT, leaf thickness; LWC, leaf water content; PH, plant height; Pn, net photosynthetic rate; RDM, root dry mass; RFM, root fresh mass; RL, root length; RWC, root water content; SDM, stem dry mass; SFM, stem fresh mass; SLA, specific leaf area; SWC, stem water content. *, p < 0.05; **, p < 0.01; ***, p < 0.001.

图8 白家溪流域自然河岸带至消落带不同植物功能性状与环境因子的冗余分析(RDA)。环境变量对功能形状的解释度见附录II。1, 2, 自然河岸带; 3-5, 过渡带; 6-8, 消落带。Chl, 叶绿素含量; Hs, 土壤湿度; IT, 淹水时长; LA, 叶面积; LDM, 叶干质量; LFM, 叶鲜质量; LL, 叶长; LT, 叶厚度; LWC, 叶含水量; PH, 植株高度; pH, 酸碱度; Pn, 净光合速率; RDM, 根干质量; RFM, 根鲜质量; RL, 根长; RWC, 根含水量; SDM, 茎干质量; SFM, 茎鲜质量; SLA, 比叶面积; SWC, 茎含水量; Ts, 土壤温度; WC, 土壤含水量; WD, 淹水深度。

Fig. 8 Redundancy analysis (RDA) analysis of plant functional traits and environmental factors from natural riparian zone to water level zone in Baijiaxi Watershed. Contribution of environmental variables to functional traits is presented in Appendix II. 1, 2, natural riparian zone; 3-5, transition zone; 6-8, drawdown zone. Chl, chlorophyll content; Hs, soil humidity; IT, inundation time; LA, leaf area; LDM, leaf dry mass; LFM, leaf fresh mass; LL, leaf length; LT, leaf thickness; LWC, leaf water content; PH, plant height; Pn, net photosynthetic rate; RDM, root dry mass; RFM, root fresh mass; RL, root length; RWC, root water content; SDM, stem dry mass; SFM, stem fresh mass; SLA, specific leaf area; SWC, stem water content; Ts, soil temperature; WC, soil water content; WD, water depth.

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三峡回水扰动增强入库小流域河岸带典型草本植物功能性状的局域分化

Intensified local differentiation in plant functional traits among typical herbaceous plants by backwater disturbance within riparian zone of small watersheds in the Three Gorges Reservoir area

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