Host Selection of Ectomycorrhizal Fungi at Pinus massoniana and Castanopsis carlesii Forests
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摘要:
目的 探究森林土壤中外生菌根真菌（Ectomycorrhizal fungi, ECMF）对宿主的选择性。 方法 采用不同的土壤菌源（马尾松土壤Pinus massoniana soil, PmS和米槠土壤Castanopsis carlesii soil, CcS），分别对马尾松（Pinus massoniana, Pm）和鳞苞锥（Castanopsis uraiana, Cu）进行接种试验（Pm-PmS、Pm-CcS、Cu-PmS、Cu-CcS）。培育6个月后，采用ITS进行菌根鉴定，检测并计算不同土壤菌源下马尾松和鳞苞锥根中ECM出现的频率、侵染率、相对丰度、相对频率、丰富度和多样性，并测定苗木生长指标以及土壤理化性质。 结果 米槠土壤pH值、全磷含量、全碳含量、有效磷含量显著高于马尾松土壤，且接种苗木后幼苗的地上干重和根长均显著高于马尾松土壤。两种土壤中共检测到19个OTUs的ECMF，分别属于7科和10属，Cenococum geophilum、Rhizopogon boninensis和Tomentella sp.2为两种土壤共有。马尾松林下土壤鉴定到的13种ECMF中，能侵染马尾松的有8种，能侵染鳞苞锥的有6种。米槠林下土壤鉴定到的9种ECMF中，能侵染马尾松的有4种，能侵染鳞苞锥的有7种。C. geophilum和Sebacina sp.2均能与马尾松和鳞苞锥建立共生关系；而Hyaloscyphaceae sp.、Lactarius inconspicuous、Rh. boninensis、Rh. flavidus、Tomentella sp.1、Tomentella sp.3和Tomentellopsis submollis只侵染马尾松；Athelia sp.、Amanita sp.、L. atrofuscus、Russula minor、Russula sp.、Sebacina sp.1、Thelephora sp.1、Thelephora sp.2和Tomentella sp.4只侵染鳞苞锥。马尾松土壤的ECMF丰富度指数（IV）、Shannon多样性指数（H´）、Simpson优势度指数（D）高于米槠土壤；但马尾松土壤接种不同宿主植物后的Sorensen相似性指数（0.14）低于米槠土壤（0.36）。部分ECMF的侵染率与寄主的生理生态指标密切相关。 结论 ECM是经过长期与树种共同进化而建立的共生关系，因此马尾松土壤中的ECMF更倾向于侵染马尾松，而米槠土壤的ECMF更倾向于侵染同为壳斗科的鳞苞锥；马尾松林下土壤的ECMF相对于米槠土壤，对寄主植物选择性的更强。虽然土壤理化性质在一定程度上影响了侵染率，但是ECMF的定殖主要受宿主植物的影响。 Abstract:Objective Host selection of ectomycorrhizal fungi (ECMF) in forest soil was studied. Method In the soils at forests of Pinus massoniana (Pm) and Castanopsis carlesii (Cc), various fungi were inoculated into Pm or Castanopsis uraiana (Cu) and designated as treatments of Pm-PmS, Pm-CcS, Cu-PmS, and Cu-CcS. After cultivating the inoculated seedlings for 6 months, mycorrhizal identification on the fungi was performed by ITS. The frequency, infection rate, relative abundance, relative frequency, richness, and diversity of ECMF in the roots of Pm and Cu plants were monitored or calculated. Seedling growth indexes and soil physiochemical properties were determined. Result The Cc forest soil (CcS) showed significantly higher pH and contents of total phosphorus, total carbon, and available phosphorus as well as the seedling shoot dry weight and root length than the Pm counterpart (PmS). The 19 OTUs of ECMF detected in these soil samples belonged to 7 families and 10 genera. Of which, Cenococum geophilum, Rhizopogon boninensis, and Tomentella sp. 2 were commonly found on the two soils. Out of the 13 ECMF identified in PmS, 8 infected Pm and 6, Cu; while among the 9 ECMF identified in CcS, 4 infected Pm and 7, Cu. Both C. geophilum and Sebacina sp. 2 were symbiotic with Pm and Cu. Hyaloscyphaceae sp., Lactarius inconspicuous, Rh. boninensis, Rh. flavidus, Tomentella sp. 1, Tomentella sp. 3 and Tomentellopsis submollis infected Pm, whereas Athelia sp., Amanita sp., L. atrofuscus, Russula minor, Russula sp., Sebacina sp.1, Thelephora sp. 1, Thelephora sp. 2 and Tomentella sp. 4 infected only Cu. The ECMF richness index (IV), Shannon index (H') and Simpson index (D) of PmS were higher than those of CcS. However, the Sorensen index on the PmS planted with host plants other than Pm was 0.14, which was lower than 0.36 on the CcS planted not with Cc. The infection rates of some ECMF were closely related to the physiological and ecological properties of the host. Conclusion The symbiosis between ECMF and trees has evolved in a long process. The ECMF in the soil at a Pm forest tended to infect Pm specifically, and so did those at a Cc forest to Cc, Cu or Fagaceae plants. However, the ECMF in PmS were more selective on their host plants than those in CcS. Even though soil physiochemical properties also affected the ECMF infection, the species of host plant largely determined the fungal colonization on the land. -
表 1 苗木接种前后土壤理化性质
Table 1. Soil physicochemical property before and after seedling inoculation
时期
Period处理
TreatmentpH值
pH value全磷
TP/（g·kg−1）全钾
TK/（g·kg−1）有效磷
AP/ （mg·kg−1）有效钾
AK/ （mg·kg−1）全碳
TC/（g·kg−1）全氮
TN/（g·kg−1）移植前
PretransplantCcS 4.82±0.02a 0.85±0.03a 14.63±0.30a 32.67±0.65a 8.46±0.61a 22.37±0.47a 2.18±0.10a PmS 4.67±0.02b 0.54±0.05b 12.73±0.31a 15.38±0.71b 6.60±0.22a 17.74±0.05b 2.20±0.02a Pm移植后
After Pm transplantationCcS 4.80±0.01a 0.82±0.01a 14.63±0.12a 33.78±0.32a 8.52±0.12a 21.83±0.16a 2.12±0.02a PmS 4.61±0.01b 0.52±0.01b 12.45±0.19a 15.17±0.04b 6.06±0.10a 17.62±0.02b 2.20±0.03a Cu移植后
After Cu transplantationCcS 4.80±0.04a 0.82±0.01a 14.41±0.17a 33.86±0.29a 8.45±0.13a 22.01±0.24a 2.18±0.03a PmS 4.64±0.01b 0.52±0.01b 12.16±0.09a 15.75±0.11b 6.81±0.15a 17.61±0.02b 2.13±0.02a Pm代表马尾松；Cu代表鳞苞锥；CcS代表米槠土壤；PmS代表马尾松土壤；不同小写字母表示独立样本t检验显著性差异（P＜0.05）。下同。
Pm: P. massoniana; Cu: C. uraiana; CcS: C. carlesii forest soil; PmS: Pm forest soil. Data with different lowercase letters indicate significant difference in Student’s t-test (P＜0.05). Same for all Tables and Figs.
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表 2 不同森林土壤处理的马尾松和鳞苞锥的生长指标
Table 2. Growth indexes of Pm and Cc under various soil treatments
处理
Treatment地上干重
Shoot dry weight/g根干重
Root dry weight/g根长
Root length/cm根表面积
Root surface area/cm2根体积
Root volume/cm3根尖数
Root tips/个Pm-PmS 0.20±0.02b 0.12±0.02a 17.95±2.00b 19.20±3.24b 0.32±0.06a 380±49.1b Pm-CcS 0.62±0.05a 0.14±0.01a 36.50±2.27a 33.60±2.72a 0.38±0.03a 570±37.9a Cu-PmS 0.36±0.03b 0.34±0.02a 18.40±1.10b 53.70±3.60a 0.41±0.04a 2399±152.0a Cu-CcS 0.58±0.04a 0.41±0.03a 33.20±1.60a 55.50±3.45a 0.45±0.03a 2755±201.0a Pm-PmS代表马尾松种植在马尾松土壤；Pm-CcS代表尾松种植在米槠土壤；Cu-PmS代表鳞苞锥种植在马尾松土壤；Cu-CcS代表鳞苞锥种植在米槠土壤。下同。
Pm-PmS: Pm planted in PmS; Pm-CcS: Pm planted in CcS; Cu-PmS: Cu planted in PmS; Cu-CcS: Cu planted in CcS. Same for all Tables and Figs.
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表 3 马尾松和米槠林外生菌根真菌OTUs的鉴定
Table 3. ECMF OTUs in PmS and CcS
序号
No.OTUs 属名
GenusNCBI对比号
NCBI comparison
number序列长度/bp（相似度/%）
Sequence length/bp
(similarity/%)相对丰度(%)/相对频率(%)
Ra(%)/Rf(%)Pm-PmS
（n=10）Cu-PmS
（n=15）Pm-CcS
（n=10）Cu-CcS
（n=15）1 Athelia sp. 阿太菌属
AtheliaAB831863 570/585 (97%) — 1.99/9.09 — — 2 Amanita sp. 鹅膏属
AmanitaOK586725 525/578 (91%) — — — 5.55/3.13 3 Cenococcum geophilum 空团菌属
CenococcumKC967408 448/449 (99%) 1.25/20.69 12.50/42.42 4.53/22.99 22.83/46.88 4 Hyaloscyphaceae sp. — MT522555 476/481 (99%) 0.46/3.45 — — — 5 Lactarius inconspicuous 乳菇属
LactariusKF433004 601/645 (93%) — — 27.35/26.44 — 6 Lactarius atrofuscus MK351919 618/635 (97%) — — — 0.26/1.56 7 Rhizopogon boninensis 须腹菌属
RhizopogonMK395368 629/633 (99%) 31.78/37.93 — 9.49/18.39 — 8 Rhizopogon flavidus KP893815 745/746 (99%) 7.82/3.45 — — — 9 Russula minor 红菇属
RussulaNR_174896 593/593 (100%) — — — 5.38/1.56 10 Russula sp. MT522574 604/607 (99%) — 7.80/9.09 — — 11 Sebacina sp.1 蜡耳壳属
SebacinaLC553305 507/542 (94%) — 7.44/6.06 — — 12 Sebacina sp.2 LC553304 531/537 (99%) — — 58.63/32.18 63.42/40.63 13 Thelephora sp.1 革菌属
ThelephoraKM576617 558/593 (94%) — 41.88/15.15 — — 14 Thelephora sp.2 HE814236 372/379 (98%) — — — 0.14/3.13 15 Tomentella sp.1 棉隔菌属
TomentellaAB848650 572/578 (99%) 37.89/17.24 — — — 16 Tomentella sp.2 HE814192 533/538 (99%) 3.61/6.90 — — 2.42/3.13 17 Tomentella sp.3 MT678909 581/583 (99%) 16.00/3.45 — — — 18 Tomentella sp.4 JF273546 572/575 (99%) — 28.40/18.18 — — 19 Tomentellopsis submollis Tomentellopsis JQ711898 609/639 (95%) 1.19/6.90 — — — “—”表示未被鉴定到。下同。
“—” means not identified. Same for below.
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表 4 马尾松和米槠林下土壤中外生菌根真菌OTUs分类统计
Table 4. OTU classification of ECMF in PmS and CcS
门
Phylum纲
Class目
Order科
Family属
GenusOTUs PmS CcS 担子菌门 Basidiomycota 伞菌纲 Agaricomycetes Boleales 须腹菌科 Rhizopogonaceae 须腹菌属 Rhizopogon 2 1 革菌目 Thelephorales 革菌科 Thelephoraceae 棉隔菌属 Tomentella 4 1 Tomentellopsis 1 0 革菌属 Thelephora 1 1 蜡耳壳目 Sebacinales 蜡耳壳科 Sebacinaceae 蜡耳壳属 Sebacina 1 1 红菇目 Russulales 红菇科 Russulaceae 乳菇属 Lactarius 0 2 红菇属 Russula 1 1 阿太菌目 Atheliales 阿太菌科 Atheliaceae 阿太菌属 Athelia 1 0 伞菌目 Agaricales 鹅膏菌科 Amanitaceae 鹅膏属 Amanita 0 1 子囊菌门 Ascomycota 锤舌菌纲 Leotiomyceces 蜡钉菌目 Helotiales Hyalodyphaceae — 1 0 座囊菌纲 Dothideomycetes 贝壳菌目 Mytilinidiales 船壳菌科 Gloniaceae 空团菌属 Cenococcum 1 1
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表 5 马尾松和米槠林下土壤处理不同宿主外生菌根真菌OTUs丰富度和多样性指数
Table 5. OTU richness and diversity of ECMF in PmS and CcS of different host plants
处理
TreatmentECMF丰富度
ECMF richness多样性指数
Shannon index优势度指数
Sinpson indexPm-PmS 13 1.72 0.77 Cu-PmS 1.60 0.77 Pm-CcS 9 1.26 0.68 Cu-CcS 1.17 0.60
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表 6 马尾松和米槠林下土壤处理不同宿主中共有外生菌根真菌类群数量(上三角)和Sorensen指数(下三角)
Table 6. Number of common ECMF groups (upper triangle) and Sorensen index (lower triangle) in PmS and CcS of different host plants
处理
TreatmentPm-PmS Cu-PmS Pm-CcS Cu-CcS Pm-PmS – 1 2 2 Cu-PmS 0.14 – 1 1 Pm-CcS 0.33 0.20 – 2 Cu-CcS 0.27 0.15 0.36 –
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表 7 马尾松林下土壤侵染马尾松和鳞苞锥幼苗的外生菌根真菌组成分析
Table 7. ECMF in PmS of infected Pm and Cu seedlings
OTUs 属名
Genus检测频率
Detection frequency/%侵染率
Infection rate/%Pm（n=10） Cu（n=15） Pm（n=10） Cu（n=15） Cenococcum geophilum 空团菌属 Cenococcum 37.5 46.7 4.0 6.1 Tomentella sp.1 棉隔菌属 Tomentella 31.3 — 67.8 — Tomentella sp.2 12.5 — 41.5 — Tomentella sp.3 6.3 — 87.0 — Tomentella sp.4 — 20.0 — 30.0 Rhizopogon boninensis 须腹菌属 Rhizopogon 68.8 — 44.5 — Rhizopogon flavidus 6.3 — 38.0 — Tomentellopsis submollis Tomentellopsis 6.3 — 4.0 — Hyaloscyphaceae sp. — 6.3 — 4.0 — Thelephora sp.1 革菌属 Thelephora — 16.7 — 48.1 Sebacina sp.1 蜡耳壳属 Sebacina — 6.7 — 31.8 Athelia sp. 阿太菌属 Athelia — 10.0 — 5.4 Russula sp. 红菇属 Russula — 10.0 — 16.9
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表 8 米槠林下土壤侵染马尾松和鳞苞锥幼苗的外生菌根真菌组成分析
Table 8. ECMF in CcS of infected Pm and Cu seedlings
属名
GenusOTUs 检测频率
Detection frequency/%侵染率
Infection rate/%Pm（n=10） Cu（n=15） Pm（n=10） Cu（n=15） 空团菌属 Cenococcum Cenococcum geophilum 66.7 100.0 8.2 10.1 蜡耳壳属 Sebacina Sebacina sp.2 93.3 86.7 54.9 31.4 乳菇属 Lactarius Lactarius inconspicuous 76.7 — 32.2 — Lactarius atrofuscus — 3.3 — 1.0 须腹菌属 Rhizopogon Rhizopogon boninensis 50.0 — 18.1 — 革菌属 Thelephora Thelephora sp.2 — 6.7 — 1.0 棉隔菌属 Tomentella Tomentella sp.2 — 6.7 — 21.5 鹅膏属 Amanita Amanita sp. — 6.7 — 44.5 红菇属 Russula Russula minor — 3.3 — 65.4
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表 9 外生菌根真菌侵染率与土壤理化性质的相关性分析
Table 9. Correlation between ECMF infection rate and soil physiochemical properties
处理
TreatmentOTUs pH值
pH value全磷
TP全钾
TK有效磷
AP有效钾
AK全碳
TC全氮
TNPm-PmS Cenococcum geophilum 0.622 0.456 −0.388 0.509 0.018 0.275 −0.469 Rhizopogon boninensis −0.314 −0.215 0.166 −0.072 0.138 −0.231 −0.435 Pm-Ccs Cenococcum geophilum 0.284 −0.079 −0.125 −0.189 −0.180 0.050 0.330 Rhizopogon boninensis −0.127 −0.029 0.206 0.086 −0.089 0.199 0.077 Sebacina sp.2 −0.022 −0.094 0.118 −0.090 0.233 0.089 −0.136 Lactarius inconspicuous −0.190 0.177 −0.209 0.271 −0.256 −0.023 0.116 Cu-PmS Cenococcum geophilum −0.074 −0.023 −0.557* 0.378 0.165 0.379 −0.319 Cu-Ccs Cenococcum geophilum 0.047 0.106 −0.027 0.074 −0.088 0.004 0.234 Sebacina sp.2 0.455* −0.121 0.074 −0.134 0.077 0.154 0.232 *表示相关在P＜0.05水平显著。
* indicates significant correlation at P＜0.05.
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表 10 外生菌根真菌侵染率与苗木生长量的相关性分析
Table 10. Correlation between ECMF infection rate and seedling biomass
处理
TreatmentOTUs 地上干重
Shoot dry weight根干重
Root dry weight根长
Root length根表面积
Root surface area根体积
Root volume根尖数
Root tipsPm-PmS Cenococcum geophilum −0.099 −0.095 −0.199 −0.065 −0.150 0.150 Rhizopogon boninensis −0.068 0.006 0.028 −0.053 0.037 0.020 Pm-Ccs Cenococcum geophilum −0.321 −0.325 −0.137 −0.337 −0.367 −0.304 Rhizopogon boninensis 0.098 −0.088 −0.12 −0.015 −0.026 −0.102 Sebacina sp.2 −0.149 −0.027 −0.324 −0.043 −0.020 0.117 Lactarius inconspicuous 0.088 −0.038 0.350 0.003 0.045 −0.139 Cu-PmS Cenococcum geophilum −0.687** −0.229 −0.370 −0.226 −0.200 −0.045 Cu-Ccs Cenococcum geophilum 0.056 −0.047 −0.074 −0.020 0.064 −0.079 Sebacina sp.2 0.192 0.004 −0.275 0.152 0.253 −0.220 **表示相关在P＜0.01水平显著。
** indicates significant correlation at P＜0.01.
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