性状调节幼苗存活率对邻体竞争和环境的响应

doi:10.17521/cjpe.2023.0271

摘要/Abstract

摘要：

性状有助于更好地理解邻体竞争和环境对于幼苗存活的影响, 然而以往研究使用物种平均性状, 忽视了种内性状变异。因此, 基于吉林蛟河针阔混交林150个幼苗样方中的幼苗个体监测数据, 运用二项分布的广义线性混合效应模型比较种内性状变异和物种平均性状对幼苗存活率的影响, 探究性状如何调节幼苗对邻体竞争和环境的响应。研究结果表明种内性状变异模型和物种平均性状模型对幼苗存活率的预测能力并不一致, 考虑种内性状变异的比叶面积(SLA)模型比物种平均性状的SLA模型赤池信息量准则值和贝叶斯信息准则值更小, 解释的方差也更多, 模型的拟合度更好。但是, 叶面积(LA)、叶碳含量(LCC)和叶氮含量(LNC)的物种平均性状模型比种内性状变异模型效果更好。此外, 性状调节邻体竞争和土壤元素含量对幼苗存活的影响, 例如: 受到相同的密度制约时, LA更小的幼苗存活率更高; 土壤营养元素缺乏时, 较高的LCC会提高幼苗存活率, 反之在优良的土壤中, 幼苗存活率降低。模型比较的结果说明种内性状变异预测幼苗个体存活的能力并不绝对比物种平均性状强, 这种原因可能与林分和环境有关。另外, 加入邻体竞争和环境变量能够增强性状与幼苗存活之间的关系。尽管种内性状变异不能更好地预测幼苗存活率, 但这种基于个体性状的方法为预测幼苗动态变化提供了一个新的视角。

关键词: 种内性状变异, 负密度制约, 生境过滤, 性状距离

Abstract:

Aims The trait-based approach has been extensively utilized to evaluate the influence of neighborhood competition on seedling survival. However, previous studies used species average traits, ignoring intraspecific trait variation.

Methods Based on the monitoring of 150 seedling plots in a needleleaf and broadleaf mixed forest in Jiaohe, Jilin, we used a generalized linear mixed effect model of binomial distribution to compare the effects of intraspecific trait variation and species average traits on seedling survival rate, and explore how functional traits mediate seedling responses to neighborhood competition and abiotic environmental factors.

Important findings The intraspecific trait variation model and the species average trait model had inconsistent predictive abilities for seedling survival rate. Specifically, intraspecific variability model of specific leaf area (SLA) exhibited a smaller Akaike information criterion value and Bayesian information criterion value, a larger explanatory variance and a better fit. While species average trait model of leaf area (LA), leaf carbon content (LCC), and leaf nitrogen content (LNC) performed better. In addition, traits mediate the effects of neighborhood competition and soil nutrients on seedling survival. Seedlings with smaller LA have higher survival rates under the same density dependence. Higher LCC increases seedling survival rate in the absence of soil nutrients, while seedling survival rate decreases under fertile soil conditions. The ability of intraspecific trait variation in predicting individual survival of seedlings may not be stronger than species average trait, which may be related to forest stands and environmental factors. In addition, the presence of neighborhood competition and environmental variables can enhance the relationships between traits and seedling survival. Although intraspecific trait variation may not improve the prediction of seedling survival rate, this individual-based approach provides a new perspective for predicting seedling dynamics.

Key words: intraspecific trait variation, negative density dependence, habitat filtering, trait dissimilarity

文佳, 张新娜, 王娟, 赵秀海, 张春雨. 性状调节幼苗存活率对邻体竞争和环境的响应. 植物生态学报, 2024, 48(6): 719-729. DOI: 10.17521/cjpe.2023.0271

WEN Jia, ZHANG Xin-Na, WANG Juan, ZHAO Xiu-Hai, ZHANG Chun-Yu. Traits mediate response of seedling survival rate to neighborhood competition and abiotic environment. Chinese Journal of Plant Ecology, 2024, 48(6): 719-729. DOI: 10.17521/cjpe.2023.0271

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2023.0271

https://www.plant-ecology.com/CN/Y2024/V48/I6/719

图/表 6

表1 吉林蛟河针阔混交林中12种乔木幼苗的物种信息

Table 1 Species information of 12 tree seedling species in a needleleaf and broadleaf mixed forest in Jiaohe, Jilin

物种 Species	科 Family	属 Genus	高度 Height (cm)	平均高度 Mean height (cm)
糠椴 Tilia mandshurica	锦葵科 Malvaceae	椴属 Tilia	4.9-12.2	7.6
杉松 Abies holophylla	松科 Pinaceae	冷杉属 Abies	4.0-31.9	7.5
水曲柳 Fraxinus mandschurica	木犀科 Oleaceae	梣属 Fraxinus	5.0-49.4	14.3
裂叶榆 Ulmus laciniata	榆科 Ulmaceae	榆属 Ulmus	5.2-40.0	14.5
大果榆 Ulmus macrocarpa	榆科 Ulmaceae	榆属 Ulmus	10.9-23.6	18.6
蒙古栎 Quercus mongolica	壳斗科 Fagaceae	栎属 Quercus	8.2-24.7	17.4
紫椴 Tilia amurensis	锦葵科 Malvaceae	椴属 Tilia	2.7-28.0	7.3
红松 Pinus koraiensis	松科 Pinaceae	松属 Pinus	3.1-30.0	11.8
东北槭 Acer mandshuricum	无患子科 Sapindaceae	槭属 Acer	4.0-48.2	16.2
色木槭 Acer pictum subsp. mono	无患子科 Sapindaceae	槭属 Acer	1.9-49.0	15.5
青楷槭 Acer tegmentosum	无患子科 Sapindaceae	槭属 Acer	8.2-19.4	12.1
千金榆 Carpinus cordata	桦木科 Betulaceae	鹅耳枥属 Carpinus	3.0-11.1	5.8

表2 吉林蛟河针阔混交林中土壤理化性质在3个主成分(PC)上的载荷值

Table 2 Loading values of soil physicochemical properties on the first three principal components (PC) in needleleaf and broadleaf mixed forest in Jiaohe, Jilin

土壤理化性质 Soil physical and chemical property	PC1	PC2	PC3
土壤含水量 Soil moisture	0.618	0.003	-0.203
全氮含量 Total nitrogen content	0.607	-0.159	-0.340
全磷含量 Total phosphorus content	0.587	0.405	0.076
全钾含量 Total potassium content	-0.596	0.461	-0.040
有机质含量 Organic matter content	0.550	-0.041	-0.201
有效氮含量 Available nitrogen content	0.634	0.487	0.298
有效磷含量 Available phosphorus content	0.051	0.626	0.405
有效钾含量 Available potassium content	0.779	-0.233	0.203
土壤pH Soil pH	0.008	-0.625	0.681
变异解释 Proportion of variance (%)	30.701	16.422	15.233
累计变异解释 Cumulative proportion of variance (%)	30.701	47.124	62.357

图1 种内性状变异和物种平均性状模型中各参数的标准化系数估计值和95%置信区间。 LA, 叶面积; LCC, 叶碳含量; LNC, 叶氮含量; SLA, 比叶面积。CI, 同种密度指数; Height, 个体高度; TI, 异种性状距离指数; Trait, 物种或个体的性状; Trait:CI, 性状和同种密度指数的交互作用; Trait:TI, 性状和异种性状距离指数的交互作用。AIC, 赤池信息量准则, 一般AIC值越小, 代表模型拟合度越好; BIC, 贝叶斯信息量准则, 一般BIC值越小, 代表模型拟合度越好。R2c, 模型的固定和随机效应对因变量的解释率; R2m, 固定效应对因变量的解释率。

Fig. 1 Estimated values and 95% confidence intervals of standardized coefficients for parameters in intraspecific trait variation and species average trait models. LA, leaf area; LCC, leaf carbon content; LNC, leaf nitrogen content; SLA, specific leaf area. CI, conspecific neighbor density index; Height, individual height; TI, heterospecific neighbor trait dissimilarity index; Trait, the trait of a species or individual; Trait:CI, the interaction between a trait and the conspecific neighbor density index; Trait:TI, the interaction between trait and the heterospecific neighbor trait dissimilarity index. AIC, Akaike Information Criterion, generally, a smaller AIC value indicates a better fit; BIC, Bayesian Information Criterion, generally, a smaller BIC value indicates a better fit. R2c, fraction of the variance explained by the fixed and random effects in the model; R2m, fraction of the variance explained by the fixed effects.

图2 性状调节幼苗存活率对于同种密度和异种邻体效应的响应。 A, 在叶面积的不同分位数下, 存活率随同种密度指数(CI)的变化。B, 在比叶面积的不同分位数下, 存活率随异种邻体性状距离指数(TI)的变化。

Fig. 2 Traits mediate the response of seedling survival rate to conspecific density dependence and heterospecific neighbor effects. A, The variation of survival rate with conspecific density index (CI) at different quantile of leaf area. B, The variation of survival rate with heterospecific neighbor trait distance index (TI) at different quantile of specific leaf area.

表3 考虑种内性状变异下包含邻体与环境变量的幼苗存活模型的各参数估计值及区间

Table 3 Considering intraspecific trait variation, the parameter estimates and confidence intervals of the seedling survival model including neighborhood and environmental variables

模型参数 Model parameter	LA	SLA	LCC	LNC
Height	0.39 [-0.05, 0.84]	0.62 [0.23, 1.00]	0.53 [0.15, 0.90]	0.46 [0.12, 0.82]
TI	-0.41 [-0.76, -0.07]	-0.18 [-0.47, 0.10]	0.05 [-0.37, 0.50]	-0.28 [-0.53, -0.03]
CI	-0.05 [-0.31, 0.22]	-0.12 [-0.40, 0.16]	-0.10 [-0.37, 0.17]	-0.07 [-0.31, 0.18]
Trait	0.70 [0.06, 1.34]	0.32 [-0.08, 0.71]	-0.66 [-1.10, -0.22]	-0.20 [-0.65, 0.01]
PC1	0.51 [0.20, 0.83]	0.48 [0.16, 0.80]	0.58 [0.25, 0.90]	0.52 [0.23, 0.81]
PC2	-0.39 [-0.70, -0.08]	-0.37 [-0.69, -0.07]	-0.34 [-0.65, -0.04]	-0.35 [-0.63, -0.07]
PC3	-0.07 [-0.40, 0.25]	-0.13 [-0.45, 0.20]	-0.18 [-0.51, 0.16]	-0.08 [-0.37, 0.22]
LAI	0.07 [-0.30, 0.44]	0.06 [-0.28, 0.40]	0.19 [-0.19, 0.57]	0.08 [-0.24, 0.40]
Trait:PC1	0.05 [-0.30, 0.40]	0.33 [0.02, 0.65]	-0.47 [-0.81, -0.14]	-0.41 [-0.68, -0.14]
Trait:PC2	-0.21 [-0.62, 0.19]	-0.03 [-0.26, 0.22]	-0.01 [-0.31, 0.29]	-0.06 [-0.30, 0.17]
Trait:PC3	0.20 [-0.14, 0.54]	0.12 [-0.13, 0.37]	0.09 [-0.29, 0.46]	-0.18 [-0.45, 0.08]
Trait:LAI	0.19 [-0.28, 0.66]	0.06 [-0.21, 0.33]	-0.30 [-0.66, 0.06]	0.13 [-0.14, 0.41]
R²_m	0.23	0.19	0.30	0.22
R²_c	0.45	0.41	0.50	0.37

图3 性状调节幼苗存活率对土壤元素含量的响应。 A, 叶碳含量(LCC)与存活率之间的斜率随土壤第一主成分(PC1)的变化。B, 土壤PC1最大和最小值的情况下, LCC与存活率之间的关系。

Fig. 3 Traits mediate the response of seedling survival rate to soil element content. A, The slope of the leaf carbon content (LCC)-seedling survival rate changes along the first principle component (PC1) of soil. B, The relationship between LCC and seedling survival rate under the maximum and minimum values of PC1 of soil.

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