亚热带天然常绿阔叶林乔木树种与林下灌木树种根-叶功能性状协调性及差异

doi:10.17521/cjpe.2024.0140

摘要/Abstract

摘要：

研究叶片和细根功能性状的协调性及差异, 有助于从植物整体的角度更好地认识植物的生态策略。为此, 该研究通过对福建省万木林自然保护区天然常绿阔叶林内20种木本植物(10种乔木、10种灌木)的叶片与根性状进行测定与分析, 探讨亚热带天然常绿阔叶林乔木与林下灌木树种根叶功能性状协调性及生存策略的差异。研究发现, 叶片与1级根相似性状间, 仅叶氮浓度和根氮浓度间存在较强的相关性, 且不受系统发育影响。群落内叶性状存在一个叶经济轴和叶组织密度-叶厚度变异轴, 1级根性状存在一个合作轴(由负相关的根直径-比根长表示)和根经济轴(由负相关的根氮浓度-根组织密度表示)。根叶经济轴之间无显著相关性。乔木和灌木间仅在根系合作轴上存在显著差异, 乔木具有较大的根直径, 而灌木具有较高的比根长。除此之外, 灌木的比叶面积显著大于乔木。研究结果表明, 亚热带天然常绿阔叶林群落内叶性状与根性状呈现复杂的整合关系, 乔木和灌木间采取不同的地上和地下策略来适应群落内的生境异质性。该研究结果扩充了对局部尺度上根叶性状间协调性的认识, 有助于深入理解群落内的生态过程和物种共存机制。

关键词: 植物功能性状, 叶经济谱, 根系经济空间, 地上-地下联系, 权衡策略, 1级根, 叶片

Abstract:

Aims Studying the coordination and differences in the functional traits of leaves and fine roots can help better understand the ecological strategies of plants from a whole-plant perspective.

Methods In this study, we measured and analyzed the leaf and root traits of 20 woody species (10 trees and 10 shrubs) from the natural evergreen broadleaf forest in Wanmulin Nature Reserve, Fujian Province. We explored the coordination of root and leaf functional traits and differences in survival strategies between tree and understory shrub species in this subtropical natural evergreen broadleaf forest.

Important findings We found a strong correlation between the leaf nitrogen concentration and root nitrogen concentration, but this was observed only for similar traits of leaf and first-order root, irrespective of phylogeny. In the studied forest, there was a leaf economics spectrum and a leaf tissue density-leaf thickness variance axis, shaped by the measured leaf traits. For first-order root, we observed a cooperative axis (represented by the negative correlation between root diameter and specific root length) and a root economics spectrum (represented by the negative correlation between root nitrogen concentration and root tissue density). There was no significant correlation between root and leaf economic spectra. Significant differences were found between tree and shrub species only along the root collaboration axis, with trees having larger root diameters and shrubs having higher specific root lengths. In addition, the specific leaf area of shrub species was significantly larger than that of tree species. The results indicated that leaf and root traits are integrated into a complex relationship, with tree and shrub species adopting different aboveground and belowground strategies to adapt to the habitat heterogeneity in the studied area. Our results expand the understanding of the coordination between root and leaf traits at a local scale, and provide deeper insights into the ecological processes and species coexistence mechanisms within the community.

Key words: plant functional trait, leaf economics spectrum, root economics space, aboveground-belowground linkages, functional trade-offs, first-order root, leaf

杜英杰, 范爱连, 王雪, 闫晓俊, 陈廷廷, 贾林巧, 姜琦, 陈光水. 亚热带天然常绿阔叶林乔木树种与林下灌木树种根-叶功能性状协调性及差异. 植物生态学报, 2025, 49(4): 585-595. DOI: 10.17521/cjpe.2024.0140

DU Ying-Jie, FAN Ai-Lian, WANG Xue, YAN Xiao-Jun, CHEN Ting-Ting, JIA Lin-Qiao, JIANG Qi, CHEN Guang-Shui. Coordination and differences in root-leaf functional traits between tree species and understory shrub species in a subtropical natural evergreen broadleaf forest. Chinese Journal of Plant Ecology, 2025, 49(4): 585-595. DOI: 10.17521/cjpe.2024.0140

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2024.0140

https://www.plant-ecology.com/CN/Y2025/V49/I4/585

图/表 7

表1 万木林自然保护区所研究树种及所属科名、生长型

Table 1 Species and family names and growth types studied in Wanmulin Nature Reserve

物种 Species	科 Family	生长型 Growth form
米槠 Castanopsis carlesii	壳斗科 Fagaceae	乔木 Tree
青冈 Cyclobalanopsis glauca	壳斗科 Fagaceae	乔木 Tree
罗浮锥 Castanopsis faberi	壳斗科 Fagaceae	乔木 Tree
苦槠 Castanopsis sclerophylla	壳斗科 Fagaceae	乔木 Tree
观光木 Michelia odora	木兰科 Magnoliaceae	乔木 Tree
沉水樟 Cinnamomum micranthum	樟科 Lauraceae	乔木 Tree
刨花润楠 Machilus pauhoi	樟科 Lauraceae	乔木 Tree
天竺桂 Cinnamomum japonicum	樟科 Lauraceae	乔木 Tree
细柄蕈树 Altingia gracilipes	蕈树科 Altingiaceae	乔木 Tree
木荷 Schima superba	山茶科 Theaceae	乔木 Tree
杜茎山 Maesa japonica	报春花科 Primulaceae	灌木 Shrub
罗浮冬青 Ilex tutcheri	冬青科 Aquifoliaceae	灌木 Shrub
三花冬青 Ilex triflora	冬青科 Aquifoliaceae	灌木 Shrub
檵木 Loropetalum chinense	金缕梅科 Hamamelidaceae	灌木 Shrub
尖叶假蚊母树 Distyliopsis dunnii	金缕梅科 Hamamelidaceae	灌木 Shrub
狗骨柴 Diplospora dubia	茜草科 Rubiaceae	灌木 Shrub
娥眉鼠刺 Itea omeiensis	鼠刺科 Iteaceae	灌木 Shrub
华幌伞枫 Heteropanax chinensis	五加科 Araliaceae	灌木 Shrub
细枝柃 Eurya loquaiana	五列木科 Pentaphylacaceae	灌木 Shrub
杨桐 Adinandra milletii	五列木科 Pentaphylacaceae	灌木 Shrub

表2 亚热带天然常绿阔叶林乔木与灌木叶与1级根性状的系统发育信号

Table 2 Phylogenetic signaling of leaf and first-order root traits of tree and shrub species in a subtropical natural evergreen broadleaf forest

叶性状 Leaf trait	K	λ	根性状 Root trait	K	λ
叶厚度 Leaf thickness	0.11	0.00	根直径 Root diameter	0.69	0.93
比叶面积 Specific leaf area	0.14	0.00	比根长 Specific root length	0.39	0.79
叶组织密度 Leaf tissue density	0.14	0.00	根组织密度 Root tissue density	0.39	0.20
叶碳浓度 Leaf carbon concentration	0.19	0.33	根碳浓度 Root carbon concentration	0.13	0.00
叶氮浓度 Leaf nitrogen concentration	0.36	0.00	根氮浓度 Root nitrogen concentration	0.83	0.92

图1 亚热带天然常绿阔叶林乔木与灌木叶与1级根性状的相关网络。A, 未去除系统发育信号。B, 去除系统发育信号。LCC, 叶碳浓度; LNC, 叶氮浓度量; LT, 叶厚度; LTD, 叶组织密度; RCC, 根碳浓度; RD, 根直径; RNC, 根氮浓度; RTD, 根组织密度; SLA, 比叶面积; SRL, 比根长。节点代表植物性状。括号中的数字表示该性状与其他性状之间的联系总数。蓝边和红边分别表示正相关和负相关(p < 0.05)。

Fig. 1 Correlation networks between leaf and first-order root traits of tree and shrub species in a subtropical natural evergreen broadleaf forest. A, Without removal of phylogenetic signals. B, With removal phylogenetic signals. LCC, leaf carbon concentration; LNC, leaf nitrogen concentration; LT, leaf thickness; LTD, leaf tissue density; RCC, root carbon concentration; RD, root diameter; RNC, root nitrogen concentration; RTD, root tissue density; SLA, specific leaf area; SRL, specific root length. Nodes represent plant traits. Numbers in parentheses indicate the total number of associations between the trait and other traits. Blue and red edges indicate positive and negative correlations, respectively (p < 0.05).

图2 亚热带天然常绿阔叶林乔木与灌木叶、1级根功能性状的主成分分析(PCA)及系统发育主成分分析(pPCA)。A, 叶片性状的主成分分析。B, 1级根性状的主成分分析。C, 叶片性状的系统发育主成分分析。D, 1级根性状的系统发育主成分分析。LCC, 叶碳浓度; LNC, 叶氮浓度; LT, 叶厚度; LTD, 叶组织密度; RCC, 根碳浓度; RD, 根直径; RNC, 根氮浓度; RTD, 根组织密度; SLA, 比叶面积; SRL, 比根长。

Fig. 2 Principal component analysis (PCA) and phylogenetic principal component analysis (pPCA) of leaf and first-order root functional traits of tree and shrub species in a subtropical natural evergreen broadleaf forest. A, PCA of leaf traits. B, PCA of first-order root traits. C, pPCA of leaf traits. D, pPCA of first-order root traits. LCC, leaf carbon concentration; LNC, leaf nitrogen concentration; LT, leaf thickness; LTD, leaf tissue density; RCC, root carbon concentration; RD, root diameter; RNC, root nitrogen concentration; RTD, root tissue density; SLA, specific leaf area; SRL, specific root length.

表3 亚热带天然常绿阔叶林叶片与1级根前两主成分间的相关关系(p值)

Table 3 Correlations between the first two principal components of leaf and first-order root in a subtropical natural evergreen broadleaf forest (p value)

	叶PC1 Leaf PC1	叶PC2 Leaf PC2	叶pPC1 Leaf pPC1	叶pPC2 Leaf pPC2
根PC1 Root PC1	-0.342 (0.14)	-0.136 (0.57)
根PC2 Root PC2	0.002 (0.99)	0.437 (0.11)
根pPC1 Root pPC1			0.342 (0.14)	0.134 (0.57)
根pPC2 Root pPC2			0.004 (0.99)	0.438 (0.11)

表4 亚热带天然常绿阔叶林乔木与灌木叶与1级根性状在经济轴得分的差异

Table 4 Differences in economic axis scores for leaf and first-order root traits between tree and shrub species in a subtropical natural evergreen broadleaf forest

主成分分析 PCA	轴 Axis	F	p	系统发育主成分分析 pPCA	轴 Axis	F	p
叶片 Leaf	PC1	0.165	0.689	叶片 Leaf	pPC1	0.165	0.689
叶片 Leaf	PC2	1.422	0.249	叶片 Leaf	pPC2	1.422	0.249
细根 Fine root	PC1	12.545	0.002	细根 Fine root	pPC1	12.485	0.002
细根 Fine root	PC2	0.795	0.384	细根 Fine root	pPC2	0.824	0.376

图3 亚热带天然常绿阔叶林乔木和灌木叶片和1级根性状对比。ns, p > 0.1; †, 0.05 < p < 0.1; *, p < 0.05; **, p < 0.01; “—”为中位数, “□”表示平均值。

Fig. 3 Comparison of leaf and first-order root traits of trees and shrubs in a subtropical natural evergreen broadleaf forest. ns, p > 0.1; †, 0.05 < p < 0.1; *, p < 0.05; **, p < 0.01; “—”is median, and “□”indicates mean.

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