氮磷添加对常绿阔叶林幼树与成树光合参数的分异影响

doi:10.17520/biods.2024435

生物多样性 ›› 2025, Vol. 33 ›› Issue (4): 24435. DOI: 10.17520/biods.2024435 cstr: 32101.14.biods.2024435

• 研究报告: 植物多样性 • 上一篇下一篇

氮磷添加对常绿阔叶林幼树与成树光合参数的分异影响

李虹茹¹, 杨桦¹, 陈伏生², 孙荣喜², 叶学敏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, 南昌 330045

收稿日期:2024-10-03 接受日期:2025-01-28 出版日期:2025-04-20 发布日期:2025-05-19
通讯作者: 叶学敏
基金资助:
国家自然科学基金(32160387);江西省林业科学院青年人才培养项目(2024520802)

Differential responses of photosynthetic parameters in saplings and adult trees to nitrogen and phosphorus addition in an evergreen broad-leaved forest

Li Hongru¹, Yang Hua¹, Chen Fusheng², Sun Rongxi², Ye Xuemin¹^,^*()

1. Nanchang National Field Observation and Research Station of Urban Ecosystem, Jiangxi Academy of Forestry, Nanchang 330032, China
2. College of Forestry, Jiangxi Agricultural University, Nanchang 330045, China

Received:2024-10-03 Accepted:2025-01-28 Online:2025-04-20 Published:2025-05-19
Contact: Ye Xuemin
Supported by:
the Nation Natural Science Foundation of China(32160387);Young Talents Training Program of Jiangxi Academy of Forestry Sciences(2024520802)

摘要/Abstract

摘要： 评估氮磷输入不平衡对亚热带森林碳汇的影响是当前的热点问题。最大羧化速率(V_cmax)和最大电子传递速率(J_max)是C₃植物光合生化模型(FvCB模型)中的重要参数, 探究氮磷养分对亚热带常绿阔叶林优势树种光合生化限制的影响机制, 有助于深化对全球变化背景下生物多样性保护和管理的认识。本研究以江西九连山常绿阔叶林中6种优势树种为研究对象, 开展为期6年的氮磷添加试验, 通过测定各个处理下叶片的V_cmax、J_max、比叶质量及氮磷含量, 分析了氮磷添加下幼树和成树的光合参数和叶片性状的特征及其相关性。结果表明, 单独添加氮或磷显著改变了幼树的光合参数, 氮添加显著提高了幼树的J_max, 磷添加显著提高了幼树的V_cmax。然而, 氮磷同添对幼树光合参数无显著影响, 表现为拮抗效应。所有氮磷处理对成树叶片光合参数均无显著影响。树木在由幼树到成树的生长过程中, V_cmax和J_max增加了3倍以上。此外, 不同物种幼树V_cmax对氮添加以及J_max/V_cmax对氮磷同添响应不同, 木荷(Schima superba)和丝栗栲(Castanopsis fargesii)对氮添加更敏感, 米槠(Castanopsis carlesii)对磷添加更敏感, 而短序润楠(Machilus breviflorai)对氮磷同添更敏感。氮磷同添显著降低了幼树的比叶质量, 提高了叶片氮含量, 磷添加显著提高了成树的叶片氮含量, 然而叶片性状并不能解释光合参数的变化。这些结果证实了理论预测, 即幼树比成树更敏感, 成树可能会通过将更多的营养资源分配给自身繁殖和防御系统, 而不是通过改变光合生化限制来调节对氮磷的利用。氮磷同添对幼树光合参数的拮抗效应表明, 养分的额外供给可能导致养分分配到非光合功能组分, 从而无法被植物光合系统高效利用。综上所述, 在常绿阔叶林中, 氮磷养分的供应显著改变了植物FvCB模型参数, 对于优势树种的光合生化限制存在个体大小依赖性和物种特异性。

关键词: 长期养分添加, 原位试验, 光合生化限制, 幼树, 成树

Abstract

Aims:Assessing the impact of nitrogen-phosphorus imbalance inputs on the carbon sink capacity of subtropical forests is currently a hot topic. Maximum carboxylation rate (V_cmax) and maximum electron transport rate (J_max) are crucial parameters in the C₃ plant photosynthetic model (FvCB model), exploring the mechanisms underlying photosynthetic biochemical limitations in response to N and P deposition, which will help to improve the understanding of biodiversity conservation and management in the context of global change.

Methods:A six-year N and P addition experiment was conducted in an evergreen broad-leaved forest of Jiulianshan, Jiangxi Province. Six dominant tree species were selected to quantify V_cmax, J_max, leaf mass per area, leaf N and P content, and analyze correlations between photosynthetic parameters and leaf traits of saplings and adult trees under N and P addition.

Results:We found that addition of either N or P significantly altered photosynthetic parameters of saplings. Specifically, N addition significantly increased the J_max of saplings, while P addition significantly increased the V_cmax. However, combined N and P addition had no significant effects on V_cmax, J_max, and J_max/V_cmax, showing an antagonistic effect. In contrast, none of N and P treatments significantly affected V_cmax, J_max, and J_max/V_cmax in adult trees. During the transition from saplings to adult trees, V_cmax and J_max increased more than threefold. Furthermore, saplings exhibited species-specific responses to N and P addition. Schima superba and Castanopsis fargesii were more sensitive to N addition, Castanopsis carlesii was more sensitive to P addition, and Machilus breviflorawas more sensitive to combined N and P addition. Combined N and P addition significantly decreased the leaf mass per area (LMA) of saplings while increasing leaf nitrogen concent, whereas P addition significantly increased leaf nitrogen content in adult trees. However, leaf traits did not explain the variation in photosynthetic parameters.

Conclusions: These findings confirm the theoretical prediction that saplings were more responsive to nutrient inputs than adult trees, which may allocate more nutrient resources to reproduction and defense systems rather than altering photosynthetic biochemical limitations in response to N and P addition. The antagonistic effect of combined N and P addition on sapling photosynthetic parameters suggests that excessive nutrient supply led to nutrient allocation toward non-photosynthetic components, reducing the efficiency of the photosynthetic system. Clearly, N and P availability significantly alter the FvCB model parameters in an evergreen broad-leaved forest, with size-dependent and species-specific effects on photosynthetic biochemical limitations.

Key words: long-term nutrient additions, in-situ experiment, photosynthetic biochemical limitation, saplings, adult trees

李虹茹, 杨桦, 陈伏生, 孙荣喜, 叶学敏 (2025) 氮磷添加对常绿阔叶林幼树与成树光合参数的分异影响. 生物多样性, 33, 24435. DOI: 10.17520/biods.2024435.

Li Hongru, Yang Hua, Chen Fusheng, Sun Rongxi, Ye Xuemin (2025) Differential responses of photosynthetic parameters in saplings and adult trees to nitrogen and phosphorus addition in an evergreen broad-leaved forest. Biodiversity Science, 33, 24435. DOI: 10.17520/biods.2024435.

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链接本文: https://www.biodiversity-science.net/CN/10.17520/biods.2024435

https://www.biodiversity-science.net/CN/Y2025/V33/I4/24435

图/表 7

表1 光合气体交换测定的物种及个体数量

Table 1 Number of individuals of different species measured by photosynthetic gas exchange

物种 Species	合计 Total	CO₂响应曲线数量 Number of CO₂ response curves
物种 Species	合计 Total	对照 Control	氮添加 N addition (+N)	磷添加 P addition (+P)	氮磷同添 Combined N + P addition (N + P)
幼树 Saplings	115	34	25	22	34
木荷 Schima superba	22	7	6	3	6
绒毛润楠 Machilus velutina	18	5	5	3	5
短序润楠 Machilus breviflora	14	4	4	2	4
丝栗栲 Castanopsis fargesii	20	6	1	5	8
米槠 Castanopsis carlesii	25	7	6	5	7
甜槠 Castanopsis eyrei	16	5	3	4	4
成树 Adult trees	45	11	12	12	10
木荷 Schima superba	12	3	3	3	3
短序润楠 Machilus breviflora	11	3	3	3	2
丝栗栲 Castanopsis fargesii	12	3	3	3	3
米槠 Castanopsis carlesii	10	2	3	3	2
所有个体 All trees	160	45	37	34	44

图1 表层土(0-10 cm)理化性质和林冠直接辐射对氮磷添加的响应。不同小写字母表示4种处理间差异显著(P < 0.05)。

Fig. 1 Response of physicochemical properties of surface soil (0-10 cm) and canopy direct radiation to N and P addition. +N, N addition; +P, P addition; N + P, Combined N + P addition. Lowercase letters indicate significant differences (P < 0.05) among the four treatments.

图2 氮磷添加对常绿阔叶林6种优势树种幼树Vcmax、Jmax和Jmax/Vcmax的影响。箱线图中, 中线代表中位数, 上下限分别表示第25和第75百分位数, 上下须线为1.5倍的四分位距(即第25和75百分位数之间的距离), 延伸出的点是离群值。图中S、N、P分别表示物种、氮添加、磷添加的效应, * P < 0.05; ** P < 0.01; *** P < 0.001。不同小写字母表示4种处理间差异显著(P < 0.05)。

Fig. 2 Effects of N and P addition on maximum carboxylation rate (Vcmax), maximum electron transport rate (Jmax), and Jmax/Vcmax in saplings of six dominant tree species in an evergreen broad-leaved forest. The line within the boxplot depicts the median, while the upper and lower bounds of the box signify the 25th and 75th percentiles, respectively. The whiskers extend to 1.5 times the interquartile range, the range between the 25th and 75th percentiles, with points beyond them considered outliers. S, N and P represent the effects of species, nitrogen addition and phosphorus addition, respectively. * P < 0.05; ** P < 0.01; *** P < 0.001. +N, N addition; +P, P addition; N + P, Combined N + P addition. Lowercase letters indicate significant differences (P < 0.05) among the four treatments.

图3 氮磷添加对常绿阔叶林4种优势树种成树Vcmax、Jmax和Jmax/Vcmax的影响。箱线图中, 中线代表中位数, 上下限分别表示第25和第75百分位数, 上下须线为1.5倍的四分位距(即第25和75百分位数之间的距离), 延伸出的点是离群值。

Fig. 3 Effects of N and P addition on maximum carboxylation rate (Vcmax), maximum electron transport rate (Jmax), and Jmax/Vcmax in adult trees of four dominant tree species in an evergreen broad-leaved forest. The line within the boxplot depicts the median, while the upper and lower bounds of the box signify the 25th and 75th percentiles, respectively. The whiskers extend to 1.5 times the interquartile range, the range between the 25th and 75th percentiles, with points beyond them considered outliers. +N, N addition; +P, P addition; N + P, Combined N + P addition.

图4 常绿阔叶林幼树和成树Vcmax和Jmax的相关性

Fig. 4 Correlation between maximum carboxylation rate (Vcmax) and maximum electron transport rate (Jmax) of saplings and adult trees in an evergreen broad-leaved forest.

图5 氮磷添加对常绿阔叶林优势树种幼树和成树比叶质量和叶片氮磷含量的影响。箱线图中, 中线代表中位数, 上下限分别表示第25和第75百分位数, 上下须线为1.5倍的四分位距(即第25和75百分位数之间的距离), 延伸出的点是离群值。图中S、N、P分别表示物种、氮添加、磷添加的效应, * P < 0.05; ** P < 0.01; *** P < 0.001。不同小写字母表示4种处理间差异显著(P < 0.05)。

Fig. 5 Effects of N and P addition on leaf mass per area (LMA), leaf N content (LNC), and leaf P content (LPC) of saplings and adult trees in an evergreen broad-leaved forest. The line within the boxplot depicts the median, while the upper and lower bounds of the box signify the 25th and 75th percentiles, respectively. The whiskers extend to 1.5 times the interquartile range, the range between the 25th and 75th percentiles, with points beyond them considered outliers. S, N and P represent the effects of species, nitrogen addition and phosphorus addition, respectively. * P < 0.05; ** P < 0.01; *** P < 0.001. +N, N addition; +P, P addition; N + P, Combined N + P addition. Lowercase letters indicate significant differences (P < 0.05) among the four treatments.

表2 氮磷添加下优势树种(幼树和成树)叶片性状和光合参数相关性分析。ns P > 0.05。

Table 2 Correlation between leaf traits and photosynthetic parameters of dominant tree species (both saplings and adult trees) in an evergreen broad-leaved forest treated with N and P addition. ns P > 0.05.

叶片性状 Leaf traits		最大羧化速率 Maximum carboxylation rate (V_cmax)	最大电子传递速率 Maximum electron transport rate (J_max)	J_max/V_cmax
比叶质量 Leaf mass per area (LMA)	P	0.157^ns	0.730^ns	0.827^ns
比叶质量 Leaf mass per area (LMA)	R²	0.09	-0.07	-0.08
叶片氮含量 Leaf N content (LNC)	P	0.248^ns	0.23^ns	0.167^ns
叶片氮含量 Leaf N content (LNC)	R²	0.047	0.09	0.08
叶片磷含量 Leaf P content (LPC)	P	0.94^ns	0.28^ns	0.45^ns
叶片磷含量 Leaf P content (LPC)	R²	-0.08	0.021	-0.03

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氮磷添加对常绿阔叶林幼树与成树光合参数的分异影响

Differential responses of photosynthetic parameters in saplings and adult trees to nitrogen and phosphorus addition in an evergreen broad-leaved forest

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