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

doi:10.17521/cjpe.2024.0281

Abstract

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

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[J]. Chin J Plant Ecol, 2025, 49(11): 1907-1918.

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Figures/Tables 7

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

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

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.

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.

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

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.

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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