磷添加对热带云雾林幼苗及幼树功能性状的影响

doi:10.17521/cjpe.2025.0361

植物生态学报 ›› 2026, Vol. 50 ›› Issue (4): 917-928.DOI: 10.17521/cjpe.2025.0361 cstr: 32100.14.cjpe.2025.0361

磷添加对热带云雾林幼苗及幼树功能性状的影响

林琳琳¹^,³, 肖劲祥¹^,³, 黄深深¹^,³, 赵杨梅²^,³, 张德旭²^,³, 程毅康²^,³^,^*(), 龙文兴²^,³^,^*()

¹ 海南大学热带农林学院(农业农村学院、乡村振兴学院), 海南儋州 571737
² 海南大学生态学院, 海口 570228
³ 海南热带雨林国家公园五指山定位观测研究站, 海南五指山 572200

收稿日期:2025-10-08 接受日期:2026-02-25 出版日期:2026-04-20 发布日期:2026-06-29
通讯作者: *程毅康, ykcheng2019@hotmail.com;
龙文兴, oklong@hainanu.edu.cn
基金资助:
国家自然科学基金(32171772);国家自然科学基金(31660163)

Effects of phosphorus addition on functional traits across seedlings and saplings in a tropical cloud forest

LIN Lin-Lin¹^,³, XIAO Jin-Xiang¹^,³, HUANG Shen-Shen¹^,³, ZHAO Yang-Mei²^,³, ZHANG De-Xu²^,³, CHENG Yi-Kang²^,³^,^*(), LONG Wen-Xing²^,³^,^*()

¹ School of Tropical Agriculture and Forestry (School of Agriculture and Rural Affairs, School of Rural Revitalization), Hainan University, Danzhou, Hainan 571737, China
² School of Ecology, Hainan University, Haikou 570228, China
³ Wuzhishan Positioning Observation and Research Station of National Park of Hainan Tropical Rainforest, Wuzhishan, Hainan 572200, China

Received:2025-10-08 Accepted:2026-02-25 Online:2026-04-20 Published:2026-06-29
Contact: *CHENG Yi-Kang, ykcheng2019@hotmail.com;
LONG Wen-Xing, oklong@hainanu.edu.cn
Supported by:
National Natural Science Foundation of China(32171772);National Natural Science Foundation of China(31660163)

摘要/Abstract

摘要： 植物功能性状是指与植物定植、存活、生长和死亡等密切相关的属性, 可独立或协同调节植物生长策略以响应环境变化。土壤磷含量是热带森林生态系统限制因子, 可能影响不同生长阶段植物的资源利用策略。该研究以海南热带雨林国家公园霸王岭片区热带云雾林的幼苗和幼树为研究对象, 设置低、中、高浓度磷添加处理, 测定植物叶片厚度(LT)、叶干物质含量(LDMC)、比叶面积(SLA)、叶片氮含量(LN)、叶片磷含量(LP)和枝条密度(BD), 分析磷添加条件下幼苗和幼树的功能性状响应及其关联性变化。结果表明: (1)植物生长阶段及磷浓度显著影响植物功能性状。在对照处理下, 相对于幼树, 幼苗表现为资源获取型策略; 随磷浓度升高, 幼苗叶片表现为高SLA、LP和低LT、LDMC, 而幼树性状变化不显著; 主成分1 (PC1)轴两端分别表现出植物的资源获取与保守型策略, 幼苗相对于幼树表现出资源获取型策略。(2)幼苗与幼树的BD-LDMC、SLA-LN、SLA-LP和LN-LP显著正关联, BD-SLA、LDMC-SLA、LDMC-LN、LDMC-LP和SLA-LT显著负关联, 表明植物功能性状间普遍存在协同与权衡关系。对照处理条件下, BD-LDMC、BD-SLA、LDMC-LN和SLA-LN的斜率在幼苗和幼树之间均无显著差异, LDMC-LP、SLA-LP和LN-LP的斜率在幼苗和幼树之间均存在显著差异, 而在磷添加处理下则相反, 说明磷添加改变了幼苗与幼树的养分分配策略。幼苗在低中磷浓度下倾向投资于资源获取有关的结构, 呈现获取型策略; 幼树在低磷条件下表现出有限的资源获取型策略调整, 对磷添加的响应相对保守。综上, 热带云雾林植物在不同生长阶段对磷添加浓度的响应存在差异, 有助于理解大气磷沉降背景下植物的资源利用及森林更新。

关键词: 磷添加, 生长阶段, 功能性状, 性状-性状关联性, 热带云雾林

Abstract:

Aims Plant functional traits reflect key strategies of resource acquisition, utilization and conservation, and play a critical role in shaping plant responses to environmental change. Soil phosphorus (P) content is a major limiting nutrient in tropical forest ecosystems, and changes in its availability may alter plant resource-use strategies at different growth stages. This study aimed to examine the responses of functional traits and their interrelationships to P addition in seedlings and saplings in a tropical cloud forest.

Methods Seedlings and saplings from the Bawangling area of National Park of Hainan Tropical Rainforest were selected as this study materials. Three P addition treatments (low, medium and high) were established. The leaf thickness (LT), leaf dry matter content (LDMC), specific leaf area (SLA), leaf nitrogen contents (LN), leaf phosphorus contents (LP) and branch density (BD) of the plants were measured. Functional traits responses and trait-trait relationships under different P addition levels were analyzed for both seedlings and saplings.

Important findings (1) The growth stage of plants and P concentration significantly influenced the functional traits of plants. Under the CK treatment, compared with saplings, seedlings exhibited a more resource acquisitive strategy. With increasing P concentration, seedlings showed higher SLA, LP and low LT, LDMC, whereas saplings exhibited no significant trait changes. The two ends of the principal component 1 (PC1) axis represented the resource-acquisitive and resource-conservative strategies, respectively, with seedlings positioned toward the acquisitive end relative to saplings. (2) BD-LDMC, SLA-LN, SLA-LP, and LN-LP of seedlings and saplings were significantly positively correlated, and BD-SLA, LDMC-SLA, LDMC-LN, LDMC-LP, and SLA-LT were significantly negatively correlated, indicating that there were generally synergies and trade-offs among plant functional traits. Under the CK treatment, the slopes of BD-LDMC, BD-SLA, LDMC-LN and SLA-LN were not significantly different between seedlings and saplings, while the slopes of LDMC-LP, SLA-LP and LN-LP were significantly different between seedlings and saplings, which was opposite under P addition treatment. This indicated that P addition changed the nutrient allocation strategy between seedlings and saplings. In the low and medium P concentration conditions, the seedlings tended to invest in resource acquisition-related structures, presenting an acquisitive strategy; while in the low P condition, the saplings showed a limited resource acquisitive strategy adjustment, and their response to P addition was relatively conservative. In conclusion, responses to P addition concentrations vary between growth stages in tropical cloud forest plants. These findings improve our understanding of plant resource-use strategies and forest regeneration dynamics under increasing atmospheric P deposition.

Key words: phosphorus addition, growth stage, functional trait, trait-trait relationship, tropical cloud forest

林琳琳, 肖劲祥, 黄深深, 赵杨梅, 张德旭, 程毅康, 龙文兴. 磷添加对热带云雾林幼苗及幼树功能性状的影响. 植物生态学报, 2026, 50(4): 917-928. DOI: 10.17521/cjpe.2025.0361

LIN Lin-Lin, XIAO Jin-Xiang, HUANG Shen-Shen, ZHAO Yang-Mei, ZHANG De-Xu, CHENG Yi-Kang, LONG Wen-Xing. Effects of phosphorus addition on functional traits across seedlings and saplings in a tropical cloud forest. Chinese Journal of Plant Ecology, 2026, 50(4): 917-928. DOI: 10.17521/cjpe.2025.0361

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2025.0361

https://www.plant-ecology.com/CN/Y2026/V50/I4/917

图/表 6

图1 热带云雾林磷添加实验样地分布图。磷添加水平(g·m-2·a-1): CK, 0; P1, 1.0; P2, 2.0; P3, 4.0; P4, 8.0; P5, 16.0。

Fig. 1 Distribution map of sample plots for phosphorus addition in a tropical cloud forest. Phosphorus addition level (g·m-2·a-1): CK, 0; P1, 1.0; P2, 2.0; P3, 4.0; P4, 8.0; P5, 16.0.

表1 磷添加与生长阶段对热带云雾林植物功能性状的影响

Table 1 Influence of phosphorus addition and growth stage on plant functional traits in a tropical cloud forest

固定效应 Fixed effect	BD		LT		SLA		LDMC		LN		LP
固定效应 Fixed effect	F	p	F	p	F	p	F	p	F	p	F	p
磷浓度 Phosphorus concentration (P)	2.38	0.07	3.30	0.02	11.52	<0.01	6.27	<0.01	0.38	0.77	38.02	<0.01
生长阶段 Growth stage (G)	20.61	<0.01	28.27	<0.01	48.73	<0.01	7.02	<0.01	0.23	0.63	10.15	<0.01
P × G	0.53	0.66	4.43	<0.01	3.22	0.02	0.60	0.61	2.58	<0.05	0.40	0.75

图2 不同磷添加水平对热带云雾林不同生长阶段植物功能性状的影响(平均值±标准差)。不同大写字母表示相同磷浓度不同生长阶段之间差异显著(p < 0.05), 不同小写字母表示相同生长阶段不同磷浓度之间差异显著(p < 0.05)。

Fig. 2 Influence of different phosphorus addition levels on plant functional traits at different growth stages in a tropical cloud forest (mean ± SD). Different uppercase letters indicate significant differences between different growth stages at the same phosphorus concentration (p < 0.05), while different lowercase letters indicate significant differences between different phosphorus concentrations at the same growth stage (p < 0.05). CK, control treatment; LoP, low phosphorus concentration addition treatment; MP, medium phosphorus concentration addition treatment; HP, high phosphorus concentration addition treatment.

图3 功能性状的方差分解分析。BD, 枝条密度; LDMC, 叶干物质含量; LN, 叶片氮含量; LP, 叶片磷含量; LT, 叶片厚度; SLA, 比叶面积。R2c, 固定效应加随机效应解释模型的拟合优度; R2m, 固定效应解释模型的拟合优度。

Fig. 3 Variance decomposition analysis of functional traits. BD, branch density; LDMC, leaf dry matter content; LN, leaf nitrogen content; LP, leaf phosphorus content; LT, leaf thickness; SLA, specific leaf area. R2c, fixed effects and random effects explanation of goodness of fit of the model; R2m, fixed effects explanation of goodness of fit of the model.

图4 不同生长阶段植物功能性状的主成分分析。A、B、C分别是幼苗、幼树和两个生长阶段的植物功能性状。BD, 枝条密度; LDMC, 叶干物质含量; LN, 叶片氮含量; LP, 叶片磷含量; LT, 叶片厚度; SLA, 比叶面积。

Fig. 4 Principal component analysis of plant functional traits at different growth stages. A, B and C are respectively the functional traits of seedlings, sapling and plants at the two growth stages. CK, control treatment; LoP, low phosphorus concentration addition treatment; MP, medium phosphorus concentration addition treatment; HP, high phosphorus concentration addition treatment. BD, branch density; LDMC, leaf dry matter content; LN, leaf nitrogen content; LP, leaf phosphorus content; LT, leaf thickness; SLA, specific leaf area.

表2 不同磷添加水平和生长阶段下的枝条与叶片功能性状间关联性

Table 2 Correlation between the functional traits of branches and leaves under different phosphorus addition levels and growth stages

Y	X	磷浓度 Phosphorus concentration	幼苗 Seeding		幼树 Sapling		Y	X	磷浓度 Phosphorus concentration	幼苗 Seeding		幼树 Sapling
Y	X	磷浓度 Phosphorus concentration	p	斜率 Slope	p	斜率 Slope	Y	X	磷浓度 Phosphorus concentration	p	斜率 Slope	p	斜率 Slope
BD	LDMC	对照 CK	0.16	1.15^Aa	<0.01	1.02^Aa	LDMC	LP	对照 CK	0.09	-0.34^Aa	<0.01	-0.58^Bab
		低磷 LoP	<0.01	1.55^Aa	<0.01	0.91^Ba			低磷 LoP	0.01	-0.56^Ab	<0.01	-0.66^Ab
		中磷 MP	<0.01	1.54^Aa	<0.01	1.09^Ba			中磷 MP	0.03	-0.61^Ab	<0.01	-0.49^Aab
		高磷 HP	<0.01	1.53^Aa	<0.01	0.96^Ba			高磷 HP	0.01	-0.33^Aa	0.03	-0.41^Aa
BD	SLA	对照 CK	0.22	-0.97^Aa	0.02	-0.71^Aab	SLA	LT	对照 CK	<0.01	-1.22^Aa	<0.01	-1.24^Aa
		低磷 LoP	<0.01	-0.95^Ba	<0.01	-0.57^Aa			低磷 LoP	0.01	-1.63^Aa	<0.01	-1.70^Ab
		中磷 MP	0.03	-1.41^Bb	<0.01	-0.80^Ab			中磷 MP	<0.01	-1.35^Aa	<0.01	-1.28^Aab
		高磷 HP	0.01	-1.24^Bab	0.03	-0.73^Aab			高磷 HP	<0.01	-1.18^Aa	<0.01	-1.24^Aab
BD	LT	对照 CK	0.22	-1.25^Ba	0.47	0.88^Aa	SLA	LN	对照 CK	<0.01	0.78^Aa	<0.01	0.89^Ab
		低磷 LoP	0.26	-1.64^Bab	0.51	0.97^Aa			低磷 LoP	0.01	0.97^Aa	<0.01	1.35^Aa
		中磷 MP	0.40	-1.77^Bb	0.99	1.02^Aa			中磷 MP	0.01	1.00^Aa	<0.01	0.93^Ab
		高磷 HP	0.56	-1.42^Bab	0.88	-0.91^Aa			高磷 HP	<0.01	0.75^Ba	<0.01	1.14^Aab
BD	LN	对照 CK	0.24	-0.70^Aa	0.09	-0.63^Aa	SLA	LP	对照 CK	0.07	0.44^Bb	<0.01	0.86^Aab
		低磷 LoP	0.36	-1.01^Aab	<0.01	-0.70^Aa			低磷 LoP	0.03	0.81^Aa	0.02	1.06^Aa
		中磷 MP	0.06	-1.36^Bb	<0.01	-0.77^Aa			中磷 MP	0.01	0.57^Aab	<0.01	0.66^Abc
		高磷 HP	0.02	-0.93^Aab	0.06	-0.78^Aa			高磷 HP	0.02	0.37^Ab	<0.01	0.56^Ac
BD	LP	对照 CK	0.82	0.37^Aa	0.09	-0.60^Bb	LT	LN	对照 CK	0.33	-0.64^Aa	<0.01	-0.70^Aa
		低磷 LoP	0.11	-0.82^Ab	<0.01	-0.55^Aab			低磷 LoP	0.18	-0.74^Aa	0.11	-0.74^Aa
		中磷 MP	0.03	-0.78^Ab	<0.01	-0.55^Aab			中磷 MP	0.78	0.89^Aa	<0.05	-0.76^Aa
		高磷 HP	0.24	-0.46^Aa	0.44	-0.38^Aa			高磷 HP	0.86	-0.76^Aa	0.02	-0.88^Aa
LDMC	SLA	对照 CK	<0.01	-0.84^Ab	<0.01	-0.69^Aa	LT	LP	对照 CK	0.14	-0.37^Aa	0.02	-0.68^Bb
		低磷 LoP	<0.01	-0.63^Aa	<0.01	-0.63^Aa			低磷 LoP	0.46	-0.61^Ab	0.81	-0.58^Aab
		中磷 MP	<0.01	-0.87^Ab	<0.01	-0.73^Aa			中磷 MP	0.83	-0.51^Aab	0.17	-0.54^Aab
		高磷 HP	<0.01	-0.81^Aab	<0.01	-0.76^Aa			高磷 HP	0.46	0.37^Aab	0.17	-0.43^Aa
LDMC	LT	对照 CK	0.39	-1.03^Aa	0.54	-0.86^Aa	LN	LP	对照 CK	0.01	0.57^Bb	<0.01	0.97^Aa
		低磷 LoP	0.76	-1.03^Aa	0.43	1.07^Aa			低磷 LoP	<0.01	0.83^Aa	<0.01	0.79^Ab
		中磷 MP	0.52	-1.17^Aa	0.28	-0.93^Aa			中磷 MP	<0.01	0.57^Ab	<0.01	0.71^Ab
		高磷 HP	0.78	-0.96^Aa	0.70	-0.94^Aa			高磷 HP	<0.01	0.49^Ab	<0.01	0.49^Ac
LDMC	LN	对照 CK	<0.01	-0.60^Aa	<0.01	-0.60^Aa
		低磷 LoP	0.01	-0.67^Aa	<0.01	-0.83^Ab
		中磷 MP	0.01	-1.06^Bb	<0.01	-0.69^Aab
		高磷 HP	<0.01	-0.66^Aa	<0.01	-0.85^Aab

表2 不同磷添加水平和生长阶段下的枝条与叶片功能性状间关联性

Table 2 Correlation between the functional traits of branches and leaves under different phosphorus addition levels and growth stages

Y	X	磷浓度 Phosphorus concentration	幼苗 Seeding		幼树 Sapling		Y	X	磷浓度 Phosphorus concentration	幼苗 Seeding		幼树 Sapling
Y	X	磷浓度 Phosphorus concentration	p	斜率 Slope	p	斜率 Slope	Y	X	磷浓度 Phosphorus concentration	p	斜率 Slope	p	斜率 Slope
BD	LDMC	对照 CK	0.16	1.15^Aa	<0.01	1.02^Aa	LDMC	LP	对照 CK	0.09	-0.34^Aa	<0.01	-0.58^Bab
		低磷 LoP	<0.01	1.55^Aa	<0.01	0.91^Ba			低磷 LoP	0.01	-0.56^Ab	<0.01	-0.66^Ab
		中磷 MP	<0.01	1.54^Aa	<0.01	1.09^Ba			中磷 MP	0.03	-0.61^Ab	<0.01	-0.49^Aab
		高磷 HP	<0.01	1.53^Aa	<0.01	0.96^Ba			高磷 HP	0.01	-0.33^Aa	0.03	-0.41^Aa
BD	SLA	对照 CK	0.22	-0.97^Aa	0.02	-0.71^Aab	SLA	LT	对照 CK	<0.01	-1.22^Aa	<0.01	-1.24^Aa
		低磷 LoP	<0.01	-0.95^Ba	<0.01	-0.57^Aa			低磷 LoP	0.01	-1.63^Aa	<0.01	-1.70^Ab
		中磷 MP	0.03	-1.41^Bb	<0.01	-0.80^Ab			中磷 MP	<0.01	-1.35^Aa	<0.01	-1.28^Aab
		高磷 HP	0.01	-1.24^Bab	0.03	-0.73^Aab			高磷 HP	<0.01	-1.18^Aa	<0.01	-1.24^Aab
BD	LT	对照 CK	0.22	-1.25^Ba	0.47	0.88^Aa	SLA	LN	对照 CK	<0.01	0.78^Aa	<0.01	0.89^Ab
		低磷 LoP	0.26	-1.64^Bab	0.51	0.97^Aa			低磷 LoP	0.01	0.97^Aa	<0.01	1.35^Aa
		中磷 MP	0.40	-1.77^Bb	0.99	1.02^Aa			中磷 MP	0.01	1.00^Aa	<0.01	0.93^Ab
		高磷 HP	0.56	-1.42^Bab	0.88	-0.91^Aa			高磷 HP	<0.01	0.75^Ba	<0.01	1.14^Aab
BD	LN	对照 CK	0.24	-0.70^Aa	0.09	-0.63^Aa	SLA	LP	对照 CK	0.07	0.44^Bb	<0.01	0.86^Aab
		低磷 LoP	0.36	-1.01^Aab	<0.01	-0.70^Aa			低磷 LoP	0.03	0.81^Aa	0.02	1.06^Aa
		中磷 MP	0.06	-1.36^Bb	<0.01	-0.77^Aa			中磷 MP	0.01	0.57^Aab	<0.01	0.66^Abc
		高磷 HP	0.02	-0.93^Aab	0.06	-0.78^Aa			高磷 HP	0.02	0.37^Ab	<0.01	0.56^Ac
BD	LP	对照 CK	0.82	0.37^Aa	0.09	-0.60^Bb	LT	LN	对照 CK	0.33	-0.64^Aa	<0.01	-0.70^Aa
		低磷 LoP	0.11	-0.82^Ab	<0.01	-0.55^Aab			低磷 LoP	0.18	-0.74^Aa	0.11	-0.74^Aa
		中磷 MP	0.03	-0.78^Ab	<0.01	-0.55^Aab			中磷 MP	0.78	0.89^Aa	<0.05	-0.76^Aa
		高磷 HP	0.24	-0.46^Aa	0.44	-0.38^Aa			高磷 HP	0.86	-0.76^Aa	0.02	-0.88^Aa
LDMC	SLA	对照 CK	<0.01	-0.84^Ab	<0.01	-0.69^Aa	LT	LP	对照 CK	0.14	-0.37^Aa	0.02	-0.68^Bb
		低磷 LoP	<0.01	-0.63^Aa	<0.01	-0.63^Aa			低磷 LoP	0.46	-0.61^Ab	0.81	-0.58^Aab
		中磷 MP	<0.01	-0.87^Ab	<0.01	-0.73^Aa			中磷 MP	0.83	-0.51^Aab	0.17	-0.54^Aab
		高磷 HP	<0.01	-0.81^Aab	<0.01	-0.76^Aa			高磷 HP	0.46	0.37^Aab	0.17	-0.43^Aa
LDMC	LT	对照 CK	0.39	-1.03^Aa	0.54	-0.86^Aa	LN	LP	对照 CK	0.01	0.57^Bb	<0.01	0.97^Aa
		低磷 LoP	0.76	-1.03^Aa	0.43	1.07^Aa			低磷 LoP	<0.01	0.83^Aa	<0.01	0.79^Ab
		中磷 MP	0.52	-1.17^Aa	0.28	-0.93^Aa			中磷 MP	<0.01	0.57^Ab	<0.01	0.71^Ab
		高磷 HP	0.78	-0.96^Aa	0.70	-0.94^Aa			高磷 HP	<0.01	0.49^Ab	<0.01	0.49^Ac
LDMC	LN	对照 CK	<0.01	-0.60^Aa	<0.01	-0.60^Aa
		低磷 LoP	0.01	-0.67^Aa	<0.01	-0.83^Ab
		中磷 MP	0.01	-1.06^Bb	<0.01	-0.69^Aab
		高磷 HP	<0.01	-0.66^Aa	<0.01	-0.85^Aab

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磷添加对热带云雾林幼苗及幼树功能性状的影响

Effects of phosphorus addition on functional traits across seedlings and saplings in a tropical cloud forest

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