氮磷添加对常绿阔叶林幼树叶片植食损伤的影响及调控机制

doi:10.17521/cjpe.2025.0248

植物生态学报 ›› 2026, Vol. 50 ›› Issue (2): 256-267.DOI: 10.17521/cjpe.2025.0248 cstr: 32100.14.cjpe.2025.0248

氮磷添加对常绿阔叶林幼树叶片植食损伤的影响及调控机制

叶学敏¹, 高伟¹, 唐星林¹, 陈伏生², 孙荣喜², 罗坤水¹^,^*()

¹ 江西省林业科学院, 南昌城市生态系统国家定位观测研究站, 南昌 330032
² 江西农业大学林学院, 亚热带森林资源培育江西省重点实验室, 南昌 330045

收稿日期:2025-06-30 接受日期:2025-12-13 出版日期:2026-02-28 发布日期:2026-04-01
通讯作者: *罗坤水 (526492282@qq.com)
基金资助:
江西省自然科学基金(20252BAC200371);江西省重点研发计划(20261BCF320043)

Effects of nitrogen and phosphorus additions on leaf herbivory damage and its underlying mechanisms in saplings in an evergreen broad-leaved forest

YE Xue-Min¹, GAO Wei¹, TANG Xing-Lin¹, CHEN Fu-Sheng², SUN Rong-Xi², LUO Kun-Shui¹^,^*()

¹ Nanchang National Field Observation and Research Station of Urban Ecosystem, Jiangxi Academy of Forestry, Nanchang 330032, China
² Jiangxi Key Laboratory of Subtropical Forest Resources Cultivation, College of Forestry, Jiangxi Agricultural University, Nanchang 330045, China

Received:2025-06-30 Accepted:2025-12-13 Online:2026-02-28 Published:2026-04-01
Contact: *LUO Kun-Shui (526492282@qq.com)
Supported by:
Jiangxi Provincial Natural Science Foundation(20252BAC200371);Jiangxi Provincial Key Research and Development Program(20261BCF320043)

摘要/Abstract

摘要：

亚热带森林中, 昆虫取食会导致叶片损伤, 但氮磷添加是否能改变幼树的抗虫能力尚不明确。该研究以江西九连山常绿阔叶林的6种优势树种幼树为研究对象, 开展为期8年的氮磷添加实验, 测定不同处理下叶片的植食损伤率、化学计量比、机械抗性及化学抗性, 分析氮磷添加对幼树抗虫性的影响。结果表明, 幼树表现出较强的抗虫能力, 叶片平均植食损伤率为5.4%, 且损伤率不受氮添加、磷添加或氮磷共同添加处理的影响, 但存在显著树种差异, 且喜光树种显著高于耐阴树种。氮添加和氮磷共同添加显著提高了叶片氮含量, 却未显著改变叶片化学计量比; 磷添加显著增加了叶片磷含量并降低了叶片碳磷比。氮磷添加对叶片撕裂力和穿透力无显著影响, 相比之下, 氮添加诱导氨基酸含量显著上升, 磷添加提高了总酚含量, 单宁含量则保持稳定。值得注意的是, 喜光树种因叶片氮含量更高而更易受害, 而耐阴树种则表现出更低的损伤率, 并在机械抗性和化学防御方面更强。综上所述, 亚热带森林幼树通过稳定的物理防御和可塑的化学防御机制, 维持对背景性植食压力的较强抵抗力, 这有助于在全球变化背景下增强群落天然更新的稳定性。

关键词: 长期养分添加, 原位试验, 叶片啃食, 次生代谢物

Abstract:

Aims Insect herbivory leads to leaf damage in subtropical forests, but it remains unclear whether nitrogen (N) and phosphorus (P) additions alter the insect resistance of saplings. This study focused on the saplings of six dominant tree species in the evergreen broad-leaved forest of Jiulian Mountain, Jiangxi Province. An eight-year N and P addition experiment was conducted to examine the effects of these nutrients on insect resistance. We measured leaf herbivory damage rate, stoichiometric ratios, mechanical resistance, and chemical resistance under different treatments.

Methods In this study, we conducted an eight-year N and P addition experiment in an evergreen broad-leaved forest in Jiulianshan, Jiangxi Province to determine effects of N and P on the saplings of six dominant tree species leaf herbivory damage, leaf stoichiometry, mechanical resistance, and chemical resistance of saplings.

Important findings Saplings exhibited strong resistance to herbivory, with an average leaf damage rate of 5.4%, which was neither affected by N addition, nor by P addition nor by both. However, significant interspecific differences were found, with light-demanding species showing significantly higher damage rates than shade-tolerant species. N addition and combined N and P addition significantly increased leaf nitrogen concentration per unit mass but did not significantly alter leaf stoichiometric ratios. P addition significantly increased leaf phosphorus concentration and decreased the carbon-to-phosphorus ratio. Both N addition and P addition showed no significant changes in leaf mechanical resistance such as force to tear and force to punch. In contrast, N addition significantly increased amino acid content, and P addition elevated total phenolics content, while tannin levels remained stable. Notably, light-demanding species exhibited higher damage due to higher leaf N concentration, whereas shade-tolerant species exhibited lower damage due to stronger mechanical resistance and chemical defense. These findings suggest that saplings in subtropical forests maintain high resistance to background insect herbivory pressure through stable physical defenses and flexible chemical defenses, potentially enhancing the ecological stability of natural regeneration within the context of global change.

Key words: long-term nutrient additions, in-situ experiment, leaf chewing, secondary metabolites

叶学敏, 高伟, 唐星林, 陈伏生, 孙荣喜, 罗坤水. 氮磷添加对常绿阔叶林幼树叶片植食损伤的影响及调控机制. 植物生态学报, 2026, 50(2): 256-267. DOI: 10.17521/cjpe.2025.0248

YE Xue-Min, GAO Wei, TANG Xing-Lin, CHEN Fu-Sheng, SUN Rong-Xi, LUO Kun-Shui. Effects of nitrogen and phosphorus additions on leaf herbivory damage and its underlying mechanisms in saplings in an evergreen broad-leaved forest. Chinese Journal of Plant Ecology, 2026, 50(2): 256-267. DOI: 10.17521/cjpe.2025.0248

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2025.0248

https://www.plant-ecology.com/CN/Y2026/V50/I2/256

图/表 9

表1 叶片植食损伤率测定的物种及个体数量

Table 1 Number of individuals of different species measured by leaf herbivorous damage rate

物种 Species	合计 Total	取样个体数量 Number of sampled individuals
物种 Species	合计 Total	对照 Control	氮添加 Nitrogen (N) addition	磷添加 Phosphorus (P) addition	氮磷共同添加 Combined N + P
木荷 Schima superba	115	30	29	29	27
短序润楠 Machilus breviflora	81	19	24	21	17
绒毛润楠 M. velutina	85	22	24	23	16
丝栗栲 Castanopsis fargesii	83	23	21	16	23
米槠 C. carlesii	76	19	16	26	15
甜槠 C. eyrei	68	20	15	18	15
幼树 Saplings	508	133	129	133	113

图1 常绿阔叶林幼树叶片植食损伤率对氮磷添加的响应(A)及种间差异(B)。不同小写字母表示不同处理或不同物种间存在显著差异(p < 0.05), ***表示喜光树种和耐阴树种存在显著差异(p < 0.001)。

Fig. 1 Responses of leaf herbivorous damage rate of saplings to nitrogen and phosphorus addition (A), interspecific differences (B) in an evergreen broad-leaved forest. Different lowercase letters indicate differences among different treatments or species (p < 0.05), *** indicate differences between light-demaning species and shade-tolerant species (p < 0.001). +N, nitrogen addition; +P, phosphorus addition; N+P, nitrogen and phosphorus combined addition.

表2 氮磷添加对常绿阔叶林中6种优势树种幼树叶片植食损伤率(LHDR)的线性混合模型(LMM)分析

Table 2 Linear mixed model (LMM) analysis for leaf herbivorous damage rate (LHDR) of six dominant tree species in an evergreen broad-leaved forest treated with nitrogen and phosphorus addition

LMM	LHDR	p	LMM	LHDR	p	LMM	LHDR	p
固定效应 Fixed effect	物种 Species	<0.001	固定效应 Fixed effect	叶片大小 Leaf size	0.002	固定效应 Fixed effect	株高 Height	0.004
	氮添加 N	0.289		氮添加 N	0.123		氮添加 N	0.265
	磷添加 P	0.217		磷添加 P	0.209		磷添加 P	0.209
	N × P	0.998		N × P	0.717		N × P	0.635
	N × species	0.052		N × leaf size	0.003		N × height	0.018
	P × species	0.724		P × leaf size	0.836		P × height	0.544
	N × P × species	0.756		N × P × leaf size	0.089		N × P × height	0.758
模型解释度 Model statistics	R²_m	0.141	模型解释度 Model statistics	R²_m	0.048	模型解释度 Model statistics	R²_m	0.034
模型解释度 Model statistics	R²_c	0.141	模型解释度 Model statistics	R²_c	0.048	模型解释度 Model statistics	R²_c	0.034

图2 常绿阔叶林幼树叶片化学计量比对氮磷添加的响应。不同小写字母表示不同处理间存在显著差异(p < 0.05)。CK, 对照; +N, 氮添加; +P, 磷添加; N+P, 氮磷共同添加。

Fig. 2 Responses of leaf stoichiometry of saplings to nitrogen and phosphorus addition in an evergreen broad-leaved forest. Different lowercase letters indicate differences among different treatments (p < 0.05). CK, control; +N, nitrogen addition; +P, phosphorus addition; N+P, nitrogen and phosphorus combined addition. Cmass, carbon content per unit mass; Nmass, nitrogen content per unit mass; Pmass, phosphorus content per unit mass.

图3 常绿阔叶林幼树比叶质量(LMA)和叶片机械抗性对氮磷添加的响应。不同小写字母表示不同处理间存在显著差异(p < 0.05)。CK, 对照; +N, 氮添加; +P, 磷添加; N+P, 氮磷共同添加。

Fig. 3 Responses of leaf mass per area (LMA) and leaf mechanical resistance of saplings to nitrogen and phosphorus addition in an evergreen broad-leaved forest. Different lowercase letters indicate differences among different treatments (p < 0.05). CK, control; +N, nitrogen addition; +P, phosphorus addition; N+P, nitrogen and phosphorus combined addition. Fp, leaf force to punch; Ft, leaf force to tear.

图4 常绿阔叶林幼树叶片化学抗性对氮磷添加的响应。不同小写字母表示不同处理间存在显著差异(p < 0.05)。CK, 对照; +N, 氮添加; +P, 磷添加; N+P, 氮磷共同添加。

Fig. 4 Responses of leaf chemical resistance of saplings to nitrogen and phosphorus addition in an evergreen broad-leaved forest. Different lowercase letters indicate differences among different treatments (p < 0.05). CK, control; +N, nitrogen addition; +P, phosphorus addition; N+P, nitrogen and phosphorus combined addition. AA, amino acid content; TA, tannin content; TPh, total phenolic content.

表3 常绿阔叶林中喜光树种和耐阴树种叶片性状的对比

Table 3 Comparison of leaf traits between light-demanding and shade-tolerant species in an evergreen broad-leaved forest

叶片性状 Leaf trait	喜光树种 Light-demanding species	耐阴树种 Shadow-tolerant species	p
叶片植食损伤率 LHDR (%)	7.13 ± 0.51	3.38 ± 0.28	^***
单位质量碳含量 C_mass (mg·g^-1)	478.19 ± 2.47	480.20 ± 3.04	0.22
单位质量氮含量 N_mass (mg·g^-1)	15.71 ± 0.32	13.79 ± 0.27	^***
单位质量磷含量 P_mass (mg·g^-1)	0.864 ± 0.026	0.824 ± 0.022	0.18
C:N	31.20 ± 0.67	35.79 ± 0.83	^***
C:P	622.2 ± 24.0	637.1 ± 20.7	0.54
N:P	19.70 ± 0.78	17.78 ± 0.57	0.13
比叶质量 LMA (g·m^-2)	89.44 ± 3.17	96.31 ± 3.36	0.06
叶片撕裂力 F_t (KN·m^-1)	0.299 ± 0.007	0.402 ± 0.009	^***
叶片穿透力 F_p (KN·m^-1)	0.198 ± 0.004	0.281 ± 0.007	^***
氨基酸含量 AA (μg·g^-1)	84.13 ± 3.66	90.83 ± 2.83	^*
总酚含量 TPh (mg·g^-1)	11.18 ± 0.79	14.68 ± 1.08	^**
单宁含量 TA (mg·g^-1)	33.21 ± 2.26	30.79 ± 1.62	0.21

图5 叶片植食损伤和叶片性状的主成分(PC)分析。前两个主成分和得分在喜光树种(灰色圆圈)和耐阴树种(绿色圆圈)中以轴表示。叶片性状缩写见表3。

Fig. 5 Principal component (PC) analysis on the leaf herbivory damage and leaf traits. The first two principal components and scores are represented along the axes for light-demanding (gray circle) and shade-tolerant (green circle) species. Abbreviations of leaf traits are provided in Table 3.

附录叶片植食损伤和叶片性状对前两个主成分的贡献 AA, 氨基酸含量; Cmass, 单位质量碳含量; Fp, 叶片穿透力; Ft, 叶片撕裂力; LHDR, 叶片植食损伤率; LMA, 比叶质量; Nmass, 单位质量氮含量; Pmass, 单位质量磷含量; PC1, 第一主成分; PC2, 第二主成分; TA, 单宁含量; TPh, 总酚含量。

Supplement Contribution of leaf herbivory damage and leaf traits to the first two principal components AA, amino acid content; Cmass, carbon content per unit mass; Fp, leaf force to punch; Ft, leaf force to tear; LHDR, leaf herbivorous damage rate; LMA, leaf mass per area; Nmass, nitrogen content per unit mass; Pmass, phosphorus content per unit mass; PC1, the first principal component; PC2, the second principal component; TA, tannin content; TPh, total phenolic content.

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氮磷添加对常绿阔叶林幼树叶片植食损伤的影响及调控机制

Effects of nitrogen and phosphorus additions on leaf herbivory damage and its underlying mechanisms in saplings in an evergreen broad-leaved forest

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