东北温带森林幼苗生长受生物量分配权衡和性状影响

doi:10.11707/j.1001-7488.LYKX20250478

摘要/Abstract

摘要：

目的: 探究东北温带森林幼苗生物量分配权衡对生长的影响及其对邻体竞争和环境因子的响应，比较生物量分配性状和器官水平性状与幼苗生长的相关性，加深对幼苗生物量分配的认识和理解，为东北温带森林的群落调控机制和动态变化规律提供理论依据。方法: 基于吉林蛟河针阔混交林130个幼苗样方个体数据，采用主成分分析确定幼苗生物量分配的权衡关系，通过线性混合效应模型构建生物量分配权衡模型、幼苗生长模型以及生物量分配性状和器官水平性状的比较模型，分析生物量分配权衡、生物量分配性状和器官水平性状对幼苗生长的影响。结果: 1）东北温带森林幼苗生物量分配权衡主要表现为2类：一是牺牲叶生物量的根生物量投资权衡，二是牺牲叶生物量和根生物量的茎生物量投资权衡。具体而言，随着土壤有机质含量减少，幼苗对根生物量的分配降低；随着林冠开阔度下降，幼苗对叶生物量的分配增加；在同种竞争和异种竞争共同作用下，幼苗会将生物量更多地分配给根或叶。2）牺牲叶生物量的根生物量投资权衡对幼苗生长具有促进作用，而林冠开阔度及土壤全磷含量和土壤全钾含量则对幼苗生长产生负向影响。3）与器官水平性状相比，生物量分配性状与幼苗生长的相关性更强，但幼苗生长率与比茎长、比叶面积、比根长等资源获取性状呈负相关。结论: 苗木生物量分配权衡对幼苗生长具有促进作用，相较于器官水平性状，生物量分配性状与幼苗生长的相关性更强，生物量分配在决定植物群落增长动态和资源吸收策略中发挥关键作用。

关键词: 生物量分配权衡, 幼苗生长, 功能性状, 温带森林, 邻体竞争

Abstract:

Objective: This study investigated the effects of seedling biomass allocation trade-offs on growth and their responses to neighborhood competition and environmental factors of temperate forests in northeast China. The correlations between biomass allocation traits and organ-level traits with seedling growth were compared. This study aims to deepen the understanding of seedling biomass allocation and provide theoretical foundations for community regulation mechanisms and dynamic patterns of temperate forests in northeast China. Method: Based on data of individual seedlings from 130 quadrats surveyed in a coniferous-broadleaved mixed forest in Jiaohe, Jilin Province, principal component analysis was used to determine the trade-offs in seedling biomass allocation. Linear mixed-effects models were employed to construct a biomass allocation trade-off model, a seedling growth model, and a comparison model for biomass allocation traits and organ-level traits, analyzing the impact of biomass allocation trade-offs as well as biomass allocation traits and organ-level traits on seedling growth. Result: 1) The biomass allocation trade-offs of temperate forest seedlings in northeast China were mainly manifested in two categories: one was the root biomass investment trade-off at the expense of leaf biomass, and the other was the stem biomass investment trade-off at the expense of both leaf and root biomass. Specifically, as soil organic matter content decreased, seedlings reduced biomass allocation to roots. As canopy openness declined, seedlings increased biomass allocation to leaves. Under the combined effects of conspecific and heterospecific competition, seedlings allocated more biomass to either roots or leaves. 2) The root biomass investment trade-off at the expense of leaf biomass promoted seedling growth, while canopy openness, soil total phosphorus content, and soil total potassium content had negative effects on seedling growth. 3) Compared with organ-level traits, biomass allocation traits were more correlated with seedling growth, but the seedling growth rate was negatively correlated with resource acquisition traits such as specific stem length, specific leaf area and specific root length. Conclusion: The trade-offs in seedling biomass allocation promote seedling growth. Compared with organ-level traits, biomass allocation traits show stronger correlations with seedling growth. These findings indicate that biomass allocation plays a crucial role in shaping plant community growth dynamics and resource acquisition strategies.

Key words: biomass allocation trade-offs, seedling growth, functional traits, temperate forests, neighbor competition

中图分类号:

S718.5

邓敏,何怀江,吴相菊,张新娜,张春雨,赵秀海. 东北温带森林幼苗生长受生物量分配权衡和性状影响[J]. 林业科学, 2026, 62(3): 48-60.

Min Deng,Huaijiang He,Xiangju Wu,Xinna Zhang,Chunyu Zhang,Xiuhai Zhao. Effects of Biomass Allocation Trade-Offs and Traits of Temperate Forests in Northeast China on Seedling Growth[J]. Scientia Silvae Sinicae, 2026, 62(3): 48-60.

图/表 14

表1

表2

图1

图2

表3

图3

表4

包含交互作用的生物量分配权衡第二主成分轴（PC2）模型结果①"

变量Variables	估计值Estimates	标准误Standard error (SE)	t值t value	P值P value
截距Intercept	0.005	0.192	0.025	0.980
高度Height	–0.126	0.059	–2.148	0.032^*
同种邻域指数Conspecific neighborhood index	–0.078	0.083	–0.940	0.348
异种邻域指数Heterospecific neighborhood index	–0.066	0.063	–1.055	0.294
土壤全氮含量Soil total nitrogen content	0.122	0.080	1.533	0.132
土壤全磷含量Soil total phosphorus content	–0.105	0.160	–0.658	0.513
土壤全钾含量Soil total potassium content	0.048	0.193	0.249	0.804
土壤有机质含量Soil organic matter content	–0.057	0.079	–0.723	0.473
叶面积指数Leaf area index	0.070	0.062	1.144	0.257
同种邻域指数和异种邻域指数交互作用 Interaction between conspecific and heterospecific neighborhood indices	–0.137	0.076	–1.803	0.072
土壤全氮含量和土壤全磷含量交互作用 Interaction between soil total nitrogen content and soil total phosphorus content	0.023	0.091	0.247	0.805
土壤全氮含量和土壤全钾含量交互作用 Interaction between soil total nitrogen content and soil total potassium content	0.293	0.116	2.518	0.017^*
土壤全氮含量和土壤有机质含量交互作用 Interaction between soil total nitrogen content and soil organic matter content	0.033	0.063	0.522	0.603
土壤全氮含量和叶面积指数交互作用 Interaction between soil total nitrogen content and leaf area index	–0.057	0.075	–0.764	0.447
土壤全磷含量和土壤全钾含量交互作用 Interaction between soil total phosphorus content and soil total potassium content	–0.030	0.199	–0.148	0.883
土壤全磷含量和土壤有机质含量交互作用 Interaction between soil total phosphorus content and soil organic matter content	0.111	0.100	1.106	0.272
土壤全磷含量和叶面积指数交互作用 Interaction between soil total phosphorus content and leaf area index	–0.043	0.095	–0.455	0.650
土壤全钾含量和土壤有机质含量交互作用 Interaction between soil total potassium content and soil organic matter content	–0.028	0.125	–0.226	0.823
土壤全钾含量和叶面积指数交互作用 Interaction between soil total potassium content and leaf area index	–0.006	0.103	–0.056	0.955
土壤有机质含量和叶面积指数交互作用 Interaction between soil organic matter content and leaf area index	–0.050	0.064	–0.782	0.436

表4

图4

表5

图5

生长模型各参数估计值和95%置信区间实心圆表示显著（P<0.05），空心圆表示不显著（P>0.05）。 Solid circles indicate significant （P<0.05）， and hollow circles indicate insignificant （P>0.05）. PC1：生物量分配权衡第一主成分轴The first principal component axis of biomass allocation trade-offs；PC2：生物量分配权衡第二主成分轴The second principal component axis of biomass allocation trade-offs；H：幼苗高度Seedling height；Acon：同种邻域指数Conspecific neighborhood index；Ahet：异种邻域指数Heterospecific neighborhood index；TN：土壤全氮含量Soil total nitrogen content；TP：土壤全磷含量Soil total phosphorus content；TK：土壤全钾含量Soil total potassium content；OM：土壤有机质含量Soil organic matter content；LAI：叶面积指数Leaf area index；Acon×Ahet：同种邻域指数与异种邻域指数交互作用Interaction between conspecific and heterospecific neighborhood indices；TN×TP：土壤全氮含量与土壤全磷含量交互作用 Interaction between soil total nitrogen content and soil total phosphorus content；TN×TK：土壤全氮含量与土壤全钾含量交互作用Interaction between soil total nitrogen content and soil total potassium content；TN×OM：土壤全氮含量与土壤有机质含量交互作用Interaction between soil total nitrogen content and soil organic matter content；TN×LAI：土壤全氮含量与叶面积指数交互作用Interaction between soil total nitrogen content and leaf area index；TP×TK：土壤全磷含量与土壤全钾含量交互作用Interaction between soil total phosphorus content and soil total potassium content；TP×OM：土壤全磷含量与土壤有机质含量交互作用Interaction between soil total phosphorus content and soil organic matter content；TP×LAI：土壤全磷含量与叶面积指数交互作用Interaction between soil total phosphorus content and leaf area index；TK×OM：土壤全钾含量与土壤有机质含量交互作用Interaction between soil total potassium content and soil organic matter content；TK×LAI：土壤全钾含量与叶面积指数交互作用Interaction between soil total potassium content and leaf area index；OM×LAI：土壤有机质含量与叶面积指数交互作用Interaction between soil organic matter content and leaf area index."

图5

图6

图7

图8

表6

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树种 Tree species	科 Family	平均高 Mean height/cm
白牛槭 Acer mandshuricum	槭树科Aceraceae	9.2
春榆Ulmus davidiana	榆科Ulmaceae	6.5
大果榆Ulmus macrocarpa	榆科Ulmaceae	12.3
红松Pinus koraiensis	松科Pinaceae	7.8
糠椴Tilia mandshurica	锦葵科Malvaceae	6.4
蒙古栎Quercus mongolica	壳斗科Fagaceae	12.7
拧筋槭Acer triflorum	无患子科Sapindaceae	12.1
千金榆Carpinus cordata	桦木科Betulaceae	5.5
青楷槭Acer tegmentosum	无患子科Sapindaceae	6.9
色木槭Acer pictum	无患子科Sapindaceae	9.3
水曲柳Fraxinus mandschurica	木樨科Oleaceae	8.0
杉松Abies holophylla	松科Pinaceae	7.0
紫椴Tilia amurensis	锦葵科Malvaceae	6.3

性状类型 Trait types	功能性状 Functional traits	缩写 Abbreviation	定义 Definition
生物量分配性状	叶质量分数	LMF	叶干质量与总干质量的比值	g·g^?1
Biomass allocation traits	Leaf mass fraction	LMF	The ratio of leaf dry mass to total dry mass	g·g^?1
Biomass allocation traits	茎质量分数	SMF	茎干质量与总干质量的比值	g·g^?1
	Stem mass fraction	SMF	The ratio of stem dry mass to total dry mass	g·g^?1
	根质量分数	RMF	根干质量与总干质量的比值	g·g^?1
	Root mass fraction	RMF	The ratio of root dry mass to total dry mass	g·g^?1
	比茎长	SSL	茎长与茎干质量的比值	cm·g^?1
	Specific stem length	SSL	The ratio of stem length to stem dry mass	cm·g^?1
器官水平性状（叶）	叶面积	LA	单叶叶片表面积	cm²
Organ-level traits (leaf)	Leaf area	LA	Single leaf surface area	cm²
Organ-level traits (leaf)	比叶面积	SLA	单位干质量的叶面积	cm²·g^?1
	Specific leaf area	SLA	Leaf surface area per dry mass	cm²·g^?1
	叶片厚度	LT	单叶叶片厚度	mm
	Leaf thickness	LT	Single leaf thickness	mm
	叶碳含量	LCC	单位叶片质量的碳含量	%
	Leaf carbon content	LCC	Carbon content per leaf mass	%
	叶氮含量	LNC	单位叶片质量的氮含量	%
	Leaf nitrogen content	LNC	Nitrogen content per leaf mass	%
器官水平性状（根）	比根长	SRL	根长与根干质量的比值	cm·g^?1
Organ-level traits (root)	Specific root length	SRL	The ratio of root length to root dry mass	cm·g^?1
Organ-level traits (root)	比根面积	SRA	单位干质量的根表面积	cm²·g^?1
	Specific root area	SRA	Root surface area per dry mass	cm²·g^?1
	根平均直径	RAD	根的平均直径	mm
	Root average diameter	RAD	Root mean diameter	mm
	根组织密度	RTD	根干质量与根体积的比值	g·cm^?3
	Root tissue density	RTD	The ratio of root dry mass to root volume	g·cm^?3

模型 Models	赤池信息准则 Akaike information criterion (AIC)	边际R² Marginal R²	条件R² Conditional R²
无交互作用的PC1模型PC1 model without interaction	1203.189	0.058	0.591
有交互作用的PC1模型PC1 model with interaction	1213.139	0.078	0.590
无交互作用的PC2模型 PC2 model without interaction	1398.564	0.026	0.146
有交互作用的PC2模型PC2 model with interaction	1404.093	0.067	0.161

模型 Models	赤池信息准则Akaike information criterion (AIC)	边际R² Marginal R²	条件R² Conditional R²
无交互作用的生长模型 Growth model without interaction	1 040.058	0.470	0.577
有交互作用的生长模型 Growth model with interaction	1 036.887	0.501	0.569

变量Variables	RDA1	RDA2	贡献 Contributions （%）	F	P
同种邻域指数Conspecific neighborhood index	0.823	0.041	36.99	19.104	<0.001^***
异种邻域指数Heterospecific neighborhood index	–0.110	–0.270	3.64	1.912	0.092
土壤全氮含量Soil total nitrogen content	–0.280	–0.557	16.99	2.817	0.027^*
土壤全磷含量Soil total phosphorus content	0.100	–0.472	9.68	3.597	0.008^**
土壤全钾含量Soil total potassium content	–0.333	0.218	7.99	5.412	<0.001^***
土壤有机质含量Soil organic matter content	–0.155	–0.219	3.28	0.403	0.871
叶面积指数Leaf area index	0.066	–0.719	21.43	2.254	0.046^*