亚热带天然林林下优势灌木叶片结构对长期土壤增温的响应

doi:10.17521/cjpe.2024.0281

植物生态学报 ›› 2025, Vol. 49 ›› Issue (11): 1907-1918.DOI: 10.17521/cjpe.2024.0281 cstr: 32100.14.cjpe.2024.0281

亚热带天然林林下优势灌木叶片结构对长期土壤增温的响应

韩润宇¹^,², 陈仕东¹^,²^,^*(), 谭艺桦¹^,², 陈相标¹^,², 熊德成¹^,², 刘小飞¹^,², 胥超¹^,², 杨智杰¹^,²^,^*(), 杨玉盛¹^,²

¹福建师范大学地理科学学院, 福州 350117
²福建三明森林生态系统国家野外科学观测研究站, 福建三明 365000

收稿日期:2024-08-21 接受日期:2025-01-09 出版日期:2025-11-20 发布日期:2025-11-20
通讯作者: *陈仕东(Chen SD, sdchen@fjnu.edu.cn);
杨智杰(Yang ZJ, zhijieyang@fjnu.edu.cn)
基金资助:
国家自然科学基金(31930071);国家自然科学基金(32192433);国家自然科学基金(32271727)

Leaf structural response of dominant understory shrubs in subtropical natural forests to long-term soil warming

HAN Run-Yu¹^,², CHEN Shi-Dong¹^,²^,^*(), TAN Yi-Hua¹^,², CHEN Xiang-Biao¹^,², XIONG De-Cheng¹^,², LIU Xiao-Fei¹^,², XU Chao¹^,², YANG Zhi-Jie¹^,²^,^*(), YANG Yu-Sheng¹^,²

¹School of Geographical Sciences, Fujian Normal University, Fuzhou 350117, China
²Fujian Sanming Forest Ecosystem National Observation and Research Station, Sanming, Fujian 365000, China

Received:2024-08-21 Accepted:2025-01-09 Online:2025-11-20 Published:2025-11-20
Supported by:
National Natural Science Foundation of China(31930071);National Natural Science Foundation of China(32192433);National Natural Science Foundation of China(32271727)

摘要/Abstract

摘要：

林下植被是亚热带森林的关键组成部分, 对维持森林生态系统结构、功能及提供生态服务等方面具有重要意义。长期土壤增温可能改变叶片功能性状进而影响林下植被的物种多样性与群落结构的演变。为探究长期土壤增温对亚热带天然林林下优势灌木叶片结构的影响, 该研究基于已进行8年的天然林原位土壤增温(+4 ℃)实验, 选取增温与对照组共有的3种优势灌木——山血丹(Ardisia lindleyana)、玉叶金花(Mussaenda pubescens)和黄毛冬青(Ilex dasyphylla), 对其叶片结构进行观察。发现增温对亚热带天然林林下优势灌木叶片的气孔与解剖结构有较大影响, 包括气孔密度和叶片厚度的减小以及比叶面积的增加, 具体表现为: 大灌木(黄毛冬青)栅栏组织和海绵组织厚度显著减少, 但下表皮厚度显著增加, 可能是增温导致叶片致密表皮毛进一步增殖的结果; 半灌木(山血丹)表皮和栅栏组织厚度显著减少, 海绵组织厚度显著增加, 反映叶片提高保水能力和缓冲水分压力的适应策略; 贴地伏生灌木(玉叶金花)上表皮和海绵组织厚度均显著减少, 但栅栏组织厚度显著增加, 表明增温提高了叶片水分运输效率及耐旱性。这些结果表明增温对亚热带天然林林下植被的水分利用效率产生显著影响, 为研究林下灌木结构对全球变暖的响应机理和生物多样性保护提供了科学依据。

关键词: 增温, 叶片结构, 林下灌木, 山血丹, 玉叶金花, 黄毛冬青

Abstract:

Aims The understory vegetation is a key component of subtropical forests closely linked to the structure and functional services of forest ecosystems. Long-term soil warming may alter leaf functional traits, affecting species diversity and community structure in the understory. In this study, we aim to investigate the effects of long-term soil warming on leaf structural succession of dominant shrubs of subtropical natural forest.

Methods Based on an eight-year in situ soil warming (+4 °C) experiment, we examined the effects of long-term soil warming on the leaf structure of three dominant shrubs (Ardisia lindleyana, Mussaenda pubescens and Ilex dasyphylla) under warming and control treatments in a natural forest.

Important findings The results indicated that long-term soil warming affected the stomatal and anatomical structure of leaves, leading to a decrease in stomatal density and leaf thickness, along with an increase in specific leaf area. Specifically, the thickness of the palisade and spongy tissue in large shrubs (Ilex dasyphylla) significantly decreased, while the thickness of the lower epidermis increased, likely due to the proliferation of dense epidermis trichomes on leaves induced by soil warming. In subshrub (Ardisia lindleyana), both epidermis and palisade tissue thickness decreased, while the spongy tissue thickness increased, reflecting an adaptive strategy to enhance water retention capacity and buffer water pressure of the leaves. In ground-lying shrubs (Mussaenda pubescens), the thickness of the upper epidermis and spongy tissue decreased, while palisade tissue thickness increased, likely enhancing water transport efficiency and drought tolerance of the leaves. Our results demonstrate that long-term soil warming has significantly impacted the water use efficiency of understory shrubs in subtropical forest. These findings provide scientific insights into the response mechanisms of understory shrub structure to global warming, with significant contribution to biodiversity conservation.

Key words: warming, leaf structure, understory shrubs, Ardisia lindleyana, Mussaenda pubescens, Ilex dasyphylla

韩润宇, 陈仕东, 谭艺桦, 陈相标, 熊德成, 刘小飞, 胥超, 杨智杰, 杨玉盛. 亚热带天然林林下优势灌木叶片结构对长期土壤增温的响应. 植物生态学报, 2025, 49(11): 1907-1918. DOI: 10.17521/cjpe.2024.0281

HAN Run-Yu, CHEN Shi-Dong, TAN Yi-Hua, CHEN Xiang-Biao, XIONG De-Cheng, LIU Xiao-Fei, XU Chao, YANG Zhi-Jie, YANG Yu-Sheng. Leaf structural response of dominant understory shrubs in subtropical natural forests to long-term soil warming. Chinese Journal of Plant Ecology, 2025, 49(11): 1907-1918. DOI: 10.17521/cjpe.2024.0281

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2024.0281

https://www.plant-ecology.com/CN/Y2025/V49/I11/1907

图/表 7

图1 格氏栲天然林样地土壤(0-10 cm)温度和含水量的时间动态(平均值±标准误)。

Fig. 1 Temporal dynamics of soil temperature and water content at 0-10 cm depth in natural forest plot of Castanopsis kawaiensis (mean ± SE).

图2 林下植被物种垂直分布状态示意图。

Fig. 2 Schematic diagram of vertical distribution of understory shrub species.

图3 山血丹、黄毛冬青和玉叶金花的气孔导度(Gsw)、气孔密度(SD)、气孔长度(SL)差异分析(平均值±标准误, n = 15)。不同大写字母表示同种植物在增温和对照间差异显著(p < 0.05), 不同小写字母表示不同植物在相同处理下差异显著(p < 0.05)。CK, 对照; P, 物种; W, 增温; W × P, 增温和物种交互。*, p < 0.05; ns, 无显著影响。

Fig. 3 Analysis of stomatal conductance for water vapor (Gsw), stomatal density (SD) and stomatal length (SL) differences among Ardisia lindleyana, Ilex dasyphylla and Mussaenda pubescens between warming and control treatments (mean ± SE, n = 15). Different uppercase letters indicate significant differences (p < 0.05) between the warming and control treatments of the same plant species, while different lowercase letters indicate significant differences (p < 0.05) between different plants under the same treatment; CK, control; P, species; W, warming; W × P, warming and species interaction. *, p < 0.05; ns, denotes no significant effect.

图4 山血丹、黄毛冬青和玉叶金花的比叶面积(SLA)、叶片厚度(LT)、上表皮厚度(UET)、栅栏组织厚度(PT)、海绵组织厚度(ST)、下表皮厚度(LET)差异分析(平均值±标准误, n = 15)。不同大写字母表示同种植物在增温和对照组之间存在显著差异(p < 0.05), 不同小写字母表示不同植物在相同处理间存在显著差异(p < 0.05)。CK, 对照; P, 物种; W, 增温; W × P, 增温和物种交互。*, p < 0.05; ns, 无显著影响。

Fig. 4 Analysis of specific leaf area (SLA), leaf thickness (LT), upper epidermis thickness (UET), palisade tissue thickness (PT), spongy tissue thickness (ST) and lower epidermis thickness (LET) differences among Ardisia lindleyana, Ilex dasyphylla and Mussaenda pubescens between warming and control treatments (mean ± SE, n = 15). Different uppercase letters indicate significant differences (p < 0.05) between the warming and control treatments of the same plant species, while different lowercase letters indicate significant differences (p < 0.05) between different plants under the same treatment. CK, control; P, species; W, warming; W × P, warming and species interaction. *, p < 0.05; ns, denotes no significant effect.

表1 山血丹、黄毛冬青和玉叶金花的叶片气孔和解剖性状因子载荷矩阵

Table 1 Factor load matrix of leaf stomatal and anatomical traits of Ardisia lindleyana, Ilex dasyphylla and Mussaenda pubescens

指标 Index	山血丹 Ardisia lindleyana		黄毛冬青 Ilex dasyphylla		玉叶金花 Mussaenda pubescens
指标 Index	PC1	PC2	PC1	PC2	PC1	PC2
气孔长度 Stomatal length	-0.172	0.384	-0.375	0.255	0.314	0.354
气孔密度 Stomatal density	0.172	-0.399	0.048	0.664	0.168	0.590
气孔导度 Stomatal conductance	0.068	0.807	-0.318	0.472	0.043	-0.646
叶片厚度 Leaf thickness	0.432	0.116	0.424	0.078	0.457	-0.034
上表皮厚度 Upper epidermis thickness	0.433	0.077	0.168	-0.378	0.445	-0.174
栅栏组织厚度 Palisade tissue thickness	0.434	0.066	0.428	0.285	-0.451	0.148
海绵组织厚度 Spongy tissue thickness	-0.428	0.125	0.445	0.161	0.451	-0.149
下表皮厚度 Lower epidermis thickness	0.437	0.049	-0.409	-0.125	0.242	0.183
特征值 Feature value	5.201	1.267	4.600	1.460	4.681	1.475

图5 山血丹、黄毛冬青和玉叶金花气孔及解剖性状主成分(PC)分析(每个点代表一个样方)。CK, 对照; Gsw, 气孔导度; LET, 下表皮厚度; LT, 叶片厚度; PT, 栅栏组织厚度; SD, 气孔密度; SL, 气孔长度; ST, 海绵组织厚度; UET, 上表皮厚度; W, 增温。

Fig. 5 Principal component (PC) analysis of stomatal and anatomical traits of Ardisia lindleyana, Ilex dasyphylla and Mussaenda pubescens (each point represents a sample plot). CK, control; Gsw, stomatal conductance for water vapor; LET, lower epidermis thickness; LT, leaf thickness; PT, palisade tissue thickness; SD, stomatal density; SL, stomatal length; ST, spongy tissue thickness; UET, upper epidermis thickness; W, warming.

表2 环境因子与山血丹、黄毛冬青和玉叶金花各指标间的相关关系分析

Table 2 Correlation analysis between environmental factors and each index of Ardisia lindleyana, Ilex dasyphylla and Mussaenda pubescens

指标 Index	山血丹 Ardisia lindleyana		黄毛冬青 Ilex dasyphylla		玉叶金花 Mussaenda pubescens
指标 Index	T	M	T	M	T	M
气孔长度 Stomatal length	0.427	-0.364	0.701^*	-0.668^*	-0.476	0.586
气孔密度 Stomatal density	-0.404	0.338	-0.076	0.204	-0.215	0.223
气孔导度 Stomatal conductance	-0.106	0.167	0.550	-0.559	-0.399	0.222
叶片厚度 Leaf thickness	-0.951^***	0.993^***	-0.962^***	0.941^***	-0.911^***	0.975^***
上表皮厚度 Upper epidermis thickness	-0.956^***	0.996^**	-0.250	0.210	-0.916^***	0.968^***
栅栏组织厚度 Palisade tissue thickness	-0.961^***	0.995^**	-0.943^***	0.986^***	0.961^***	-0.992^***
海绵组织厚度 Spongy tissue thickness	0.951^***	-0.963^***	-0.970^***	0.995^***	-0.958^***	0.996^***
下表皮厚度 Lower epidermis thickness	-0.964^***	0.996^***	0.839^**	-0.811^**	-0.395	0.439

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亚热带天然林林下优势灌木叶片结构对长期土壤增温的响应

Leaf structural response of dominant understory shrubs in subtropical natural forests to long-term soil warming

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