缙云山4种林地山莓根际土壤真菌群落分析

doi:10.16420/j.issn.0513-353x.2025-0073

摘要/Abstract

摘要：

为合理利用我国林地资源，探寻山莓（Rubus corchorifolius）林下种植的适宜林地类型，以山莓灌木丛为参照对象，选取重庆缙云山中亚热带典型植被中的马尾松（Pinus massoniana）、楠竹（Phyllostachys ubescens）和柑橘（Citrus reticulata）3种林地的山莓根际土壤，采用高通量测序法检测其真菌多样性，并分析它们与土壤养分间的关系。结果发现，马尾松和山莓林地中山莓根际土壤真菌多样性指数间无显著差异，但都显著高于楠竹和柑橘林地；子囊菌门（Ascomycota）、担子菌门（Basidiomycota）和被孢霉门（Mortierellomycota）是山莓根际土壤的优势真菌门，它们的总相对丰度在山莓、柑橘和马尾松林地中均占90%以上，在楠竹林地中占40%以上；山莓、马尾松、柑橘和楠竹林地间的关键差异菌属明显不同，依次分别为：篮状菌属（Talaromyces）和弯颈霉属（Tolypocladium）、绿僵菌属（Metarhizium）和青霉属（Penicillium）、曲霉属（Aspergillus）和毛壳菌属（Chaetomium）、未分类的子囊菌门菌属（Unclassified_k_Ascomycota）。土壤中的有效磷和有机碳含量是影响山莓根际土壤真菌群落分布的关键养分因子。山莓根际土壤真菌的主要功能类群在马尾松和山莓林地中均为腐生真菌，而在楠竹和柑橘林地中则分别为丛枝菌根真菌和寄生真菌。综上，4种林地中，马尾松和山莓林地间的山莓根际土壤养分良好，关键差异菌属均可能具有解磷功能，且真菌的物种组成相近、丰富度和多样性都显著高于柑橘和楠竹林地，可推荐用作林下栽培山莓的适宜林地。

关键词: 山莓, 林地, 真菌多样性, 土壤养分

Abstract:

This study aims to promote the rational utilization of China’s forest land resources and to identify suitable forest types for the understory cultivation of Rubus corchorifolius. Using R. corchorifolius shrubland as a reference，its rhizosphere soil were investigated in three typical mid-subtropical forest stands i.e. Pinus massoniana，Phyllostachys ubescens，and Citrus reticulata in Jinyun Mountain，Chongqing. High-throughput sequencing was employed to analyze fungal diversity and its relationship with soil nutrients. Results showed that the fungal diversity index of rhizosphere soil was no significantly different between Pinus massoniana and R. corchorifoliu stands，but was significantly higher than that in Phyllostachys ubescens and C. reticulata stands. Ascomycota，Basidiomycota，and Mortierellomycota were the dominant fungal phyla，collectively accounting for over 90% of the totally relative abundance under R. corchorifolius，C. reticulata，and P. massoniana stands，while > 40% under Phyllostachys ubescens stand. The key differential fungal genera distinctly varied among the four stands：Talaromyces and Tolypocladium in the R. corchorifolius stand；Metarhizium and Penicillium in the Pinus massoniana stand；Aspergillus and Chaetomium in the C. reticulata stand；and Unclassified_k_Ascomycota in the Phyllostachys ubescens stand. Soil available phosphorus and organic carbon were the critical nutrient factors influencing the distribution of fungal communities in R. corchorifolius rhizosphere soil. The saprotrophic fungi were the primary functional groups under both Pinus massoniana and R. corchorifolius stands，whereas arbuscular mycorrhizal fungi and parasitic fungi were predominant under Phyllostachys ubescens and C. reticulata stands. Overall，the rhizosphere soil nutrients were favorable under Pinus massoniana and R. corchorifolius stands than under other two stands while the major key fungal genera might have the function of phosphorus solubilization. In conclusion，the Pinus massoniana and R. corchorifolius stands exhibited more favorable rhizosphere soil nutrients. Their key differential genera are likely capable of phosphorus solubilization，and they shared similar fungal species composition with significantly higher richness and diversity compared to the C. reticulata and Phyllostachys ubescens stands. Therefore，the Pinus massoniana forest stand is recommended as a suitable site for the understory cultivation of R. corchorifolius.

Key words: Rubus corchorifolius, forest stand, fungal diversity, soil nutrient

吕亚茹, 李杰, 辜夕容, 崔瑶, 胡佳. 缙云山4种林地山莓根际土壤真菌群落分析[J]. 园艺学报, 2026, 53(1): 202-218.

LÜ Yaru, LI Jie, GU Xirong, CUI Yao, HU Jia. Analysis of Fungal Community in Rhizosphere Soil of Rubus corchorifolius Under Four Forest Stands in Jinyun Mountain[J]. Acta Horticulturae Sinica, 2026, 53(1): 202-218.

图/表 12

表1 缙云山4种林地植被主要植物种类及山莓果实品质

Table 1 Dominant plant species and fruit quality of Rubus corchorifolius under four forest stands in Mount Jinyun

林地类型 Forest type	植物种类 Plant species	山莓果实品质 Rubus corchorifolius fruit quality
山莓 Rubus corchorifolius	山莓、葛藤、芭茅、火炭母 Rubus corchorifolius，Argyreia pierreana，Miscanthus floridulus，Polygonum chinense	果中偏大，果甜较酸，香味多，味浓风味佳 The fruit is medium-large in size，with a sweet and moderately acidic flavor，a rich aroma，and a full-bodied，excellent taste
马尾松 Pinus massoniana	马尾松、山莓、杉木、毛桐、接骨草 Pinus massoniana，Rubus corchorifolius，Cunninghamia lanceolata，Mallotus japonicus，Sambucus chinensis	果中偏大，果甜微酸，香味多，有特殊香气，味浓风味好 The fruit is medium-large in size，with a sweet and slightly acidic flavor，a rich and distinctive aroma，and a full-bodied，pleasant taste
楠竹 Phyllostachys ubescens	楠竹、山莓、蕨、鸢尾 Phyllostachys ubescens，Rubus corchorifolius，Pteridium aquilinum，Iris tectorum	果中偏小，果较甜微酸，香味偏淡，籽多 The fruit is medium-small in size，with a relatively sweet and slightly acidic flavor，a faint aroma，and a high seed content
柑橘 Citrus reticulata	柑橘、山莓、燕麦、苎麻 Citrus reticulate，Rubus corchorifolius，Avena sativa，Boehmeria nivea	果中偏小，果甜微酸，籽粒脆，香味较多，风味较好 The fruit is medium-small in size，with a sweet and slightly acidic flavor，crisp seeds，a pronounced aroma，and a good overall taste

表2 缙云山4种林地山莓根际土壤养分含量

Table 2 Nutrient content in rhizosphere soil of Rubus corchorifolius under 4 forest stands in Jinyun Mountain

林地类型 Forest stand	pH	有机碳/（g · kg^-1） Organic carbon （SOC）	总氮/（g · kg^-1） Total N （TN）	总磷/（g · kg^-1） Total P （TP）	总钾/（g · kg^-1） Total K（TK）
山莓Rubus corchorifolius	4.79 ± 0.04 a	12.60 ± 0.02 b	1.61 ±0.00 b	0.37 ± 0.02 b	10.80 ± 0.05 c
马尾松Pinus massoniana	4.69 ± 0.03 b	12.30 ± 0.02 c	1.96 ± 0.02 a	1.07 ± 0.03 a	19.50 ± 0.03 a
楠竹Phyllostachys ubescens	4.31 ± 0.01 c	25.60 ± 0.13 a	1.54 ± 0.02 c	0.37 ± 0.02 b	12.20 ± 0.07 b
柑橘Citrus reticulata	4.37 ± 0.01 c	8.67 ± 0.08 d	1.16 ± 0 d	0.34 ± 0.01 b	8.71 ± 0.07 d
林地类型 Forest stand	碳氮比 C/N	氮磷比 N/P	碱解氮/（mg · kg^-1）Available N（AN）	有效磷/（mg · kg^-1）Available P（AP）	速效钾/（mg · kg^-1）Available K（AK）
山莓Rubus corchorifolius	7.79 ± 0.00 b	4.39 ± 0.27 a	124.00 ± 1.17 a	72.60 ± 0.63 a	75.80 ± 0.10 b
马尾松Pinus massoniana	6.24 ± 0.07 c	1.83 ± 0.07 c	100.00 ± 1.03 b	44.10 ± 0.09 b	133.00 ± 0.27 a
楠竹Phyllostachys ubescens	16.60 ± 0.37 a	4.16 ± 0.21 a	99.30 ± 0.06 b	24.60 ± 0.32 c	24.00 ± 0.04 d
柑橘Citrus reticulata	7.43 ± 0.09 b	3.47 ± 0.10 b	83.90 ± 1.29 c	14.20 ± 0.03 d	39.80 ± 0.01 c

图1 缙云山4种林地中山莓根际土壤真菌的稀释曲线图（A）和OTU分布Venn图（B） RC：山莓林地；CR：柑橘林地；PU：楠竹林地；PM：马尾松林地。下同

Fig. 1 Dilution curve（A）and Venn diagram（B）of OTU distribution in rhizosphere soil fungi of Rubus corchorifolius under four forest stands in Jinyun Mountain RC：Rubus corchorifolius stand；CR：Citrus reticulata stand；PU：Phyllostachys ubescen stand；PM：Pinus massoniana stand. The same below

图2 缙云山4种林地中山莓根际土壤真菌的Sobs（A）、Chao（B）、Ace（C）和香农指数（D）不同小写字母表示林分间差异显著（P < 0.05）

Fig. 2 Indexes of Sobs（A），Chao（B），Ace（C）and Shannon（D）of rhizosphere soil fungi of Rubus corchorifolius under four forest stands in Jinyun Mountain Different lowercase letters indicate significant differences among different forest stands at 0.05 level

图3 缙云山4种林地中山莓根际土壤真菌群落的主坐标分析（PCoA）

Fig. 3 Principal Co-ordinates Analysis（PCoA）of the rhizosphere soil fungal community of Rubus corchorifolius under four forest stands in Jinyun Mountain

图4 缙云山4种林地中山莓根际土壤真菌门水平分类结果（A）及显著性比较（B） A中的“*”表示经Kruskal-Wallis秩和检验多组间存在显著差异，B中的“*”表示经Wilcoxon秩和检验两两组间差异显著，P < 0.05

Fig. 4 Classification results（A）and difference comparison（B）of the rhizosphere soil fungal at the phylum level of Rubus corchorifolius under four forest stands in Jinyun Mountain The asterisk * in Fig. A indicates a significant difference among multiple groups（Kruskal-Wallis test），while in Fig. B，it signifies a significant difference between two groups（Wilcoxon test），P < 0.05 for both

图5 缙云山4种林地中山莓根际土壤真菌属水平分类结果（A）及显著性比较（B） “*”表示经Kruskal-Wallis秩和检验在多组间存在显著差异，P < 0.05

Fig. 5 Classification results（A）and difference comparison（B）of the rhizosphere soil fungal at the genus level of the Rubus corchorifolius under four forest stands in Jinyun Mountain The asterisk in Fig. 5，B indicates a significant difference among multiple groups（Kruskal-Wallis test）（P < 0.05）

图6 缙云山4种林地中山莓根际土壤真菌群落的关键差异性菌属不同颜色的节点表示微生物在不同组中显著富集，浅黄色的节点表示不同组间无显著差异。P：门；C：纲；F：科；O：目；G：属

Fig. 6 Biomarker with the rhizosphere soil of Rubus corchorifolius under four forest stands in Jinyun Mountain Nodes of different colors indicate that microorganisms are significantly enriched in different groups，while light yellow nodes indicate no significant differences between groups. P：Phylum；C：Class；F：Family；O：Order；G：Genus

图7 缙云山4种林地中山莓根际土壤真菌群落与土壤养分的典范对应分析（CCA）各养分代号详见表2，下同。图中不同颜色或形状的点表示不同环境或条件下的样本组，红色箭头表示数量型环境因子。从样品点向数量型环境因子的箭头做投影，投影点距离原点的距离代表环境因子对样本群落分布相对影响的大小，点与箭头的方向是否一致代表正、负相关性

Fig. 7 Canonical Correspondence Analysis（CCA）of the rhizosphere soil fungal community of Rubus corchorifolius with soil nutrients content under four forest stands in Jinyun Mountain Codes for nutrients refer to Table 2，and the same applies hereafter. In the diagram，samples from different environments or conditions are represented by points of varying colors or shapes. Red arrows indicate quantitative environmental variables. By projecting sample points onto the directional arrows of quantitative environmental variables，the distance from the projection point to the origin reflects the relative influence of the environmental variable on community distribution，while the directional alignment between sample points and arrows indicates positive or negative correlations

图8 缙云山4种林地中山莓根际土壤养分与优势菌属间的相关性 N/P：氮磷比；C/N：碳氮比；SOC：有机碳；AN：碱解氮；AP：有效磷；AK：速效钾；TN：总氮；TK：总钾；TP：总磷

Fig. 8 Correlation between the rhizosphere soil nutrients of Rubus corchorifolius and dominant genus under four forest stands in Jinyun Mountain N/P：Nitrogen-phosphorus ratio；C/N：Carbon-nitrogen ratio；SOC：Organic carbon；AN：Available N；AP：Available P；AK：Available K；TN：Total N；TK：Total K；TP：Total P. *：P < 0.05，**：P < 0.01，***：P < 0.001

表3 缙云山4种林地中山莓根际土壤真菌群落多样性指数与土壤养分的相关性

Table 3 Correlation between rhizosphere soil fungal community and soil nutrients of Rubus corchorifolius under four forest stands in Jinyun Mountain

指数 Index	pH	有机碳 Organic carbon（SOC）	总氮 Total N （TN）	总磷 Total P （TP）	总钾 Total K （TK）	碳氮比 C/N	氮磷比 N/P	碱解氮 Available N（AN）	有效磷Available P（AP）	速效钾Available K（AK）
Ace	0.880^**	-0.632^*	0.547	0.553	0.456	-0.799^**	-0.390	0.499	0.704^*	0.845^**
Chao	0.879^**	-0.626^*	0.543	0.546	0.450	-0.792^**	-0.380	0.506	0.707^*	0.838^**
香农指数 Shannon’s index	0.731^**	0.005	0.725^**	0.503	0.540	-0.230	-0.238	0.683^*	0.704^*	0.615^*
生态优势指数 Sobs index	0.894^**	-0.560	0.615^*	0.588^*	0.513	-0.748^**	-0.395	0.538	0.729^**	0.864^**

图9 缙云山4种林地中山莓根际土壤的真菌群落预测功能分类

Fig. 9 Functional classification of fungal communities in the rhizosphere of Rubus corchorifolius under four forest stands in Jinyun Mountain

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