林业科学 ›› 2026, Vol. 62 ›› Issue (10): 42-57.doi: 10.11707/j.1001-7488.LYKX20260402
陈思梦1,王爽1,刘倩愿1,*(
),寇亮2,杨风亭2,王辉民2
收稿日期:2026-06-26
出版日期:2026-10-10
发布日期:2026-10-16
通讯作者:
刘倩愿
E-mail:liuqianyuan333@126.com
基金资助:
Simeng Chen1,Shuang Wang1,Qianyuan Liu1,*(
),Liang Kou2,Fengting Yang2,Huimin Wang2
Received:2026-06-26
Online:2026-10-10
Published:2026-10-16
Contact:
Qianyuan Liu
E-mail:liuqianyuan333@126.com
摘要:
目的: 探究亚热带常绿阔叶林不同菌根类型的林下木本植物根叶功能性状差异,揭示菌根类型是否影响根叶性状关联以及哪些土壤因子影响丛枝菌根和外生菌根植物根叶性状变异,为理解菌根类型如何驱动植物地上?地下功能性状协同变异以及植物的环境适应策略提供理论依据。方法: 以亚热带常绿阔叶林中共存的5个丛枝菌根(AM)和5个外生菌根(ECM)树种为对象,通过测定植物叶片和根系功能性状以及土壤理化性质,分析不同菌根类型植物根叶功能性状差异,并阐明性状变异与土壤因子的关系。结果: 丛枝菌根树种具有更高的叶片含水量、比叶面积、比根长和比根面积,比叶质量、叶碳含量、根组织密度、根分枝比、分枝强度和根碳含量则显著低于外生菌根树种。丛枝菌根树种倾向于采取“低投入–快速回报”的资源获取型策略,即通过高比叶面积和低比叶质量快速获取资源,外生菌根树种则倾向于采取“高投入?缓慢回报”的资源保守型策略,具有低比叶面积和高比叶质量,资源获取速率较慢但防御能力较强。根性状变异还存在独立的自身根系构建(高比根长与比根面积)与菌根共生投资(高分枝强度与依赖菌根共生获取资源)的“自己动手–菌根投资”协作轴:丛枝菌根树种依赖高比根长和比根表面积增强自身根系对土壤养分和水分的获取能力,外生菌根树种则通过增加根系分枝促进与真菌共生增强资源获取。根叶性状相关性在2类菌根类型间差异明显:丛枝菌根树种比叶面积与根分枝强度呈显著正相关,反映其地上–地下资源获取策略的协同一致性;而外生菌根树种上述关系不显著,呈现解耦格局。土壤因子对2类菌根树种的根叶性状配置具有差异化调控作用。在低pH、低无机氮含量条件下,丛枝菌根树种表现出比叶面积和比根长增大;在高pH和低硝态氮含量条件下,外生菌根树种增大根组织密度和根分枝强度。土壤可溶性有机碳含量与丛枝菌根植物的比根长和比表面积呈显著正相关(P<0.05),与外生菌根植物的比叶面积呈显著正相关(P<0.05),而与外生菌根植物的叶碳含量呈极显著负相关(P<0.001)。结论: 菌根类型驱动根叶性状沿资源“获取型?保守型”和“自己动手–菌根投资”双轴分化,土壤pH和氮有效性是根叶性状变异的关键环境过滤器。
中图分类号:
陈思梦,王爽,刘倩愿,寇亮,杨风亭,王辉民. 亚热带常绿阔叶林不同菌根类型的林下木本植物根叶功能性状差异及影响因素[J]. 林业科学, 2026, 62(10): 42-57.
Simeng Chen,Shuang Wang,Qianyuan Liu,Liang Kou,Fengting Yang,Huimin Wang. Variations in Root and Leaf Functional Traits and Their Driving Factors in Understory Woody Plants with Different Mycorrhizal Types in Subtropical Evergreen Broad-Leaved Forests[J]. Scientia Silvae Sinicae, 2026, 62(10): 42-57.
表1
亚热带10种林下优势木本植物基本信息"
| 物种 Species | 菌根类型Mycorrhizal type | 树高 Tree height/m | 地径 Ground diameter/mm |
| 黄瑞木Adinandra millettii | 丛枝菌根Arbuscular mycorrhiza | 2.58 | 39.60 |
| 荚蒾Viburnum dilatatum | 丛枝菌根Arbuscular mycorrhiza | 2.08 | 22.40 |
| 栀子树Gardenia jasminoides | 丛枝菌根Arbuscular mycorrhiza | 1.58 | 13.08 |
| 枫香Liquidambar formosana | 丛枝菌根Arbuscular mycorrhiza | 2.60 | 28.43 |
| 盐肤木Rhus chinensis | 丛枝菌根Arbuscular mycorrhiza | 6.90 | 72.00 |
| 檵木Loropetalum chinense | 外生菌根Ectomycorrhiza | 3.55 | 42.60 |
| 青冈栎Quercus glauca | 外生菌根Ectomycorrhiza | 3.95 | 71.10 |
| 麻栎Quercus acutissima | 外生菌根Ectomycorrhiza | 4.15 | 92.98 |
| 湿地松Pinus elliottii | 外生菌根Ectomycorrhiza | 5.93 | 127.75 |
| 红锥Castanopsis hystrix | 外生菌根Ectomycorrhiza | 5.05 | 49.20 |
表2
植物叶片和根系性状的统计描述"
| 项目Items | 简写 Abbreviation | 单位 Units | 样本数量 Sample numbers | 最小值 Min. | 最大值 Max. | 均值 Mean | 标准误差 Standard error | 变异系数 Coefficient of variation |
| 叶片含水量Leaf water content | LWC | % | 40 | 31.02 | 69.94 | 53.61 | 1.36 | 0.16 |
| 比叶面积Specific leaf area | SLA | cm2·g–1 | 40 | 21.17 | 196.62 | 118.39 | 6.79 | 0.36 |
| 比叶质量Leaf mass per area | LMA | mg·cm–2 | 40 | 5.09 | 47.24 | 10.82 | 1.34 | 0.79 |
| 叶碳含量Leaf carbon content | LCC | g·g–1 | 40 | 0.45 | 0.54 | 0.50 | 0.00 | 0.04 |
| 叶氮含量Leaf nitrogen content | LNC | mg·g–1 | 40 | 11.84 | 25.03 | 17.56 | 0.52 | 0.19 |
| 叶碳氮比Leaf carbon-to-nitrogen ratio | LC∶N | — | 40 | 20.72 | 42.91 | 29.16 | 0.84 | 0.18 |
| 根直径Root diameter | RD | mm | 40 | 0.11 | 0.27 | 0.17 | 0.01 | 0.27 |
| 根组织密度Root tissue density | RTD | g·cm–3 | 40 | 0.29 | 1.11 | 0.71 | 0.04 | 0.32 |
| 比根长Specific root length | SRL | m·g–1 | 40 | 32.70 | 193.35 | 77.50 | 6.90 | 0.56 |
| 比根表面积Specific root surface area | SRA | cm2·g–1 | 40 | 201.59 | 749.41 | 369.87 | 22.36 | 0.38 |
| 根分枝比Root branching ratio | BR | — | 40 | 1.43 | 8.48 | 4.00 | 0.31 | 0.48 |
| 根分枝强度Root branching density | BI | cm–1 | 40 | 1.34 | 19.49 | 6.16 | 0.72 | 0.74 |
| 根碳含量Root carbon content | RCC | g·g–1 | 40 | 0.36 | 0.47 | 0.43 | 0.00 | 0.06 |
| 根氮含量Root nitrogen content | RNC | mg·g–1 | 40 | 7.38 | 14.91 | 10.88 | 0.30 | 0.18 |
| 根碳氮比Root carbon-to-nitrogen ratio | RC∶N | — | 40 | 25.69 | 57.87 | 40.88 | 1.29 | 0.20 |
图3
叶性状(A)和根性状(B)的主成分分析 LWC:叶片含水量Leaf water content;SLA:比叶面积Specific leaf area;LNC:叶氮含量Leaf nitrogen content;LCC:叶碳含量Leaf carbon content;LMA:比叶质量Leaf mass per area;LC∶N:叶碳氮比Leaf carbon-to-nitrogen ratio;RNC:根氮含量Root nitrogen content;SRA:比根表面积Specific root surface area;SRL:比根长Specific root length;RC∶N:根碳氮比Root carbon-to-nitrogen ratio;RCC:根碳含量Root carbon content;RTD:根组织密度Root tissue density;BI:根分枝强度Root branching density;BR:根分枝比Root branching ratio;RD:根直径Root diameter."
表3
叶性状在主成分(PC1和PC2)上的载荷矩阵①"
| 叶片性状Leaf traits | PC1 | PC2 |
| 叶片含水量Leaf water content (LWC) | ?0.019 | ?0.727 |
| 比叶面积Specific leaf area (SLA) | ?0.877 | ?0.285 |
| 比叶质量Leaf mass per area (LMA) | 0.823 | 0.314 |
| 叶碳含量Leaf carbon content (LCC) | 0.248 | 0.842 |
| 叶氮含量Leaf nitrogen content (LNC) | ?0.779 | 0.560 |
| 叶碳氮比Leaf carbon-to-nitrogen ratio (LC∶N) | 0.899 | ?0.328 |
| 特征值Eigenvalues | 2.923 | 1.840 |
| 方差解释率Ratio of variance explained (%) | 48.72 | 30.67 |
| 累积解释率Cumulative resolution rate (%) | 48.72 | 79.39 |
表4
根性状在主成分(PC1和PC2)上的载荷矩阵①"
| 根性状Root traits | PC1 | PC2 |
| 根直径Root diameter (RD) | 0.145 | ?0.656 |
| 根组织密度Root tissue density (RTD) | 0.816 | 0.231 |
| 比根长Specific root length (SRL) | ?0.753 | 0.483 |
| 比根表面积Specific root surface area (SRA) | ?0.901 | 0.281 |
| 根分枝比Root branching ratio (BR) | 0.803 | ?0.153 |
| 根分枝强度Root branching density (BI) | 0.786 | 0.104 |
| 根碳含量Root carbon content (RCC) | 0.410 | 0.432 |
| 根氮含量Root nitrogen content (RNC) | ?0.189 | ?0.810 |
| 根碳氮比Root carbon-to-nitrogen ratio (RC∶N) | 0.274 | 0.858 |
| 特征值Eigenvalues | 3.607 | 2.408 |
| 方差解释率Ratio of variance explained (%) | 40.07 | 26.76 |
| 累积解释率Cumulative resolution rate (%) | 40.07 | 66.83 |
图5
叶根性状联合主成分分析以及土壤理化性质与主成分坐标轴得分的线性回归 SLA:比叶面积Specific leaf area;SRL:比根长 Specific root length;SRA:比根表面积 Specific root surface area;LWC:叶片含水量 Leaf water content;LC∶N:叶碳氮比 Leaf carbon-to-nitrogen ratio;RNC:根氮含量 Root nitrogen content;RD:根直径 Root diameter;LMA:比叶质量 Leaf mass per area;LCC:叶碳含量 Leaf carbon content;BR:根分枝比 Root branching ratio;BI:根分枝强度 Root branching density;RTD:根组织密度 Root tissue density;RCC:根碳含量 Root carbon content;LNC:叶氮含量 Leaf nitrogen content;RC∶N:根碳氮比 Root carbon-to-nitrogen ratio."
表5
根叶性状联合在主成分上的载荷矩阵①"
| 性状Traits | PC1 | PC2 |
| 叶片含水量Leaf water content (LWC) | ?0.726 | 0.281 |
| 比叶面积Specific leaf area (SLA) | ?0.378 | ?0.643 |
| 比叶质量Leaf mass per area (LMA) | 0.367 | 0.802 |
| 叶碳含量Leaf carbon content (LCC) | 0.580 | 0.283 |
| 叶氮含量Leaf nitrogen content (LNC) | 0.250 | ?0.610 |
| 叶碳氮比Leaf carbon-to-nitrogen ratio (LC∶N) | ?0.085 | 0.709 |
| 根直径Root diameter (RD) | 0.137 | 0.521 |
| 根组织密度Root tissue density (RTD) | 0.756 | ?0.265 |
| 比根长Specific root length (SRL) | ?0.708 | ?0.222 |
| 比根表面积Specific root surface area (SRA) | ?0.852 | ?0.083 |
| 根分枝比Root branching ratio (BR) | 0.821 | 0.018 |
| 根分枝强度Root branching density (BI) | 0.826 | ?0.157 |
| 根碳含量Root carbon content (RCC) | 0.377 | ?0.438 |
| 根氮含量Root nitrogen content (RNC) | ?0.040 | 0.558 |
| 根碳氮比Root carbon-to-nitrogen ratio (RC∶N) | 0.143 | ?0.569 |
| 特征值Eigenvalues | 4.552 | 3.342 |
| 方差解释率Ratio of variance explained (%) | 30.35 | 22.28 |
| 累积解释率Cumulative resolution rate (%) | 30.35 | 52.63 |
图6
所有物种(A)、丛枝菌根(B)和外生菌根(C)物种的根叶性状和土壤因子的相关性 LWC:叶片含水量Leaf water content;SLA:比叶面积Specific leaf area;LMA:比叶质量Leaf mass per area;LCC:叶碳含量Leaf carbon content;LNC:叶氮含量Leaf nitrogen content;LC∶N:叶碳氮比Leaf carbon-to-nitrogen ratio;RD:根直径Root diameter;RTD:根组织密度Root tissue density;SRL:比根长Specific root length;SRA:比根表面积Specific root surface area;BR:根分枝比Root branching ratio;BI:根分枝强度Root branching density;RCC:根碳含量Root carbon content;RNC:根氮含量Root nitrogen content;RC∶N:根碳氮比Root carbon-to-nitrogen ratio;EC:土壤电导率Soil electrical conductivity;SWC:土壤含水量Soil water content;SOC:土壤有机碳含量Soil organic carbon content;DOC:土壤可溶性有机碳含量Soil dissolved carbon content;STN:土壤全氮含量Soil total nitrogen content;AMN:土壤铵态氮含量Soil ammonium nitrogen content;NTN:土壤硝态氮含量Soil nitrate nitrogen content;STP:土壤全磷含量Soil total phosphorus content;SC∶N:土壤碳氮比Soil carbon-to-nitrogen ratio;SC∶P:土壤碳磷比Soil carbon-to-phosphorus ratio;SN∶P:土壤氮磷比Soil nitrogen-to-phosphorus ratio. *: P<0.05; **: P<0.01; ***: P<0.001."
表6
冗余分析前的土壤因素共线性检验"
| 土壤因子Soil factors | 方差膨胀 因子VIF |
| pH | 2.698 |
| 土壤电导率Soil electrical conductivity (EC) | 2.416 |
| 土壤含水量Soil water content (SWC) | 1.290 |
| 土壤有机碳含量Soil organic carbon content (SOC) | 6.037 |
| 土壤可溶性有机碳含量Soil dissolved carbon content (DOC) | 1.544 |
| 土壤全氮含量Soil total nitrogen content (STN) | 5.894 |
| 土壤铵态氮含量Soil ammonium nitrogen content (AMN) | 2.861 |
| 土壤硝态氮含量Soil nitrate nitrogen content (NTN) | 4.444 |
| 土壤全磷含量Soil total phosphorus content (STP) | 1.603 |
图7
土壤理化性质对叶性状(A)和根性状(B)的影响的冗余分析以及相对应的土壤变量解释率(C、D) 红色箭头代表土壤理化性质,蓝色箭头分别代表叶片性状(A)和根系性状(B)。The red arrows represent soil physicochemical properties, the blue arrows represent leaf traits(A) and root traits (B), respectively. LWC:叶片含水量Leaf water content;STP:土壤全磷含量Soil total phosphorus content;DOC:土壤可溶性有机碳含量Soil dissolved carbon content;SLA:比叶面积Specific leaf area;STN:土壤全氮含量Soil total nitrogen content;SOC:土壤有机碳含量Soil organic carbon content;SWC:土壤含水量Soil water content;LNC:叶氮含量Leaf nitrogen content;NTN:土壤硝态氮含量Soil nitrate nitrogen content;EC:土壤电导率Soil electrical conductivity;LCC:叶碳含量Leaf carbon content;LMA:比叶质量Leaf mass per area;AMN:土壤铵态氮含量Soil ammonium nitrogen content;SRA:比根表面积Specific root surface area;SRL:比根长Specific root length;RNC:根氮含量Root nitrogen content;RCC:根碳含量Root carbon content;RTD:根组织密度Root tissue density;BI:根分枝强度Root branching density;BR:根分枝比Root branching ratio;RD:根直径Root diameter. *: P<0.05; **: P<0.01; ***: P<0.001."
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