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转基因玉米连续种植对土壤丛枝菌根真菌群落的影响
Effects of Continuous Planting of Transgenic Maize on Community of Arbuscular Mycorrhizal Fungi in Soil
为评估转基因玉米连续种植对土壤丛枝菌根真菌群落结构的影响，以转基因玉米DBN9936和其受体玉米DBN318为材料，采用PCR-DGGE技术和PLFA技术分析土壤AM真菌群落结构特征。结果表明：在相同的种植年份和地点下，转基因和非转基因玉米的土壤AM真菌群落结构表现出较高的相似性，其香农-威纳指数、Pielou均匀度指数和丰富度均未出现显著性差异，且优势属均为Clomus（球囊霉属）。聚类分析结果显示，同一种植地点下的2种玉米土壤中AM真菌群落的相似度大多在0.60以上，而不同地点的群落相似度均小于0.60；同时，不同种植地点的两种玉米非常清晰的分布在聚类图的上下两侧，说明土壤AM真菌群落结构受种植地点影响较大，而转基因本身对其影响较为微弱。系统发育分析结果表明，不同地点下种植的两种玉米的特有条带在系统发育树上聚类在不同的类群。土壤微生物PLFA含量结果显示，转基因玉米DBN9936的种植对土壤微生物的总PLFA含量均未产生显著影响，各微生物类群的相对丰度差异也较小；而在不同种植地点下两种玉米土壤微生物的总PLFA含量均有显著性差异，且各微生物类群的相对丰度差异较大。综上，转基因玉米DBN9936较之对应的非转基因玉米对土壤AM真菌群落结构和PLFA含量无显著影响，但不同种植地点之间存在差异。
To evaluate the effect of continuous planting of transgenic maize on the community structure of soil arbuscular mycorrhizal fungi, transgenic maize DBN9936 and its non-transgenic counterpart were used as the experimental materials, the characteristics of soil arbuscular mycorrhizal (AM) fungal community structure were analyzed using PCR-DGGE and PLFA techniques. The results indicated that there was a high similarity in the soil AM fungal community structure between transgenic and non-transgenic maize in the same planting year and location. There were no significant differences in the Shannon-Wiener index, Pielou evenness index and richness. The dominant genera consistently identified were Glomus. Cluster analysis revealed that the similarity of AM fungal communities in soils from the two maize types at the same planting location was mostly above 0.60, while the similarity between communities from different locations was below 0.60. Additionally, the clustering pattern clearly separated the two maize types from different planting locations on the upper and lower sides of the cluster diagram, indicated that the community structure of soil AM fungi was greatly affected by planting sites, while the influence of the transgenic itself was weak. Phylogenetic analysis also indicated that the specific bands of the two maize species planted in different locations clustered in different groups on the phylogenetic trees. The results of soil microbial PLFA content showed that the planting of transgenic maize DBN9936 had no significant effect on the total soil microbial PLFA content, and the differences in the relative abundance of microbial groups were small. In contrast, significant differences were observed in the total PLFA content of soil microbes between the two maize types planted at different locations, with substantial variations in the relative abundance of microbial groups. In conclusion, compared to non-transgenic maize, transgenic maize DBN9936 had no significant impact on the structure of soil AM fungal communities and PLFA content. However, notable differences were observed among different planting locations.
转基因玉米 / 丛枝菌根真菌 / PCR-DGGE / PLFA / 群落结构 {{custom_keyword}} /
transgenic maize / arbuscular mycorrhizal fungi / PCR-DGGE / PLFA / community structure {{custom_keyword}} /
表1 试验地土壤基本理化性质 |
| 试验地 | 全磷/(g/kg) | 全氮/(g/kg) | 土壤有机质/(g/kg) | pH |
|---|---|---|---|---|
| 吉林 | 0.52 | 1.02 | 9.35 | 5.16 |
| 河北 | 0.53 | 0.49 | 9.35 | 8.37 |
表2 PCR反应的引物及反应条件 |
| 巢式PCR | 引物 | 引物序列(5′-3′) | 反应条件 |
|---|---|---|---|
| 第一步 | AML1 | ATCAACTTTCGATGGTAGGATAGA | 94℃ 3 min；94℃ 30 s，50℃ 30 s， 72℃ 45 s，35个循环；72℃ 7 min |
| AML2 | GAACCCAAACACTTTGGTTTCC | ||
| 第二步 | NS31 | GC-CGCCCGCCGCGCGCGGCGGGCGGGGCGGGGGCA CGGGGGGTTGGA | 94℃ 3 min；94℃ 30 s，50℃ 30 s， 72℃ 45 s，35个循环；72℃ 7 min |
| G101 | GGGCAA GTCTGGTGCC GCC TGC TTT AAA CAC TCT A |
表3 磷脂脂肪酸类群分类 |
| 微生物类群 | 磷脂脂肪酸标记 |
|---|---|
| 细菌 | i14:0，14:0，i15:0，a15:0，15:0，i16:0，16:0，16:1ω7c，i17:0，a17:0，17:0，cy17:0，18:1ω7，i19:0，a19:0，19:0，cy19:0 |
| 革兰氏阳性细菌 | i14:0，i15:0，a15:0，i16:0，i17:0，a17:0 |
| 革兰氏阴性细菌 | 16:1ω7c，16:1ω9c，cyc17:0，17:1ω8c，18:1ω7c，cyc19:0 |
| 真菌 | 16:1ω5c，18:1ω9c，18:2ω6c，18:3ω6c |
| AM真菌 | 16:1ω5c，18:1ω7 |
| 厌氧菌 | A17:0，i17:0 |
表4 2016年土壤AM真菌DGGE条带对比结果 |
| 条带 编号 | GeneBank 登录号 | 同源性最高的菌株 | 相似度/% |
|---|---|---|---|
| 1 | JQ218148.1 | Uncultured Glomus clone | 100 |
| 2 | KX809136.1 | Uncultured Rhizophagus clone | 99 |
| 3 | KY979384.1 | Uncultured Glomus clone | 100 |
| 6 | KM365412.1 | Uncultured Glomeraceae clone | 100 |
| 7 | KC579419.1 | Uncultured Glomus clone | 99 |
| 8 | KY232438.1 | Uncultured Glomus clone | 99 |
| 9 | LN906586.1 | Uncultured Archaeospora clone | 99 |
| 10 | KU168035.1 | Uncultured Claroideoglomu | 100 |
| 12 | LT856617.1 | Uncultured Rhizophagus clone | 99 |
| 13 | LT856616.1 | Uncultured Rhizophagus clone | 100 |
| 14 | EU332725.1 | Uncultured Paraglomus clone | 98 |
| 15 | GQ140597.1 | Uncultured Glomus clone | 100 |
| 16 | KC588997.1 | Uncultured Glomus clone | 98 |
| 17 | JN559802.1 | Uncultured Glomus clone | 98 |
| 18 | MG835506.1 | Uncultured Glomus clone | 99 |
| 19 | KY979290.1 | Uncultured Glomus clone | 100 |
| 20 | KM085113.1 | Uncultured Archaeospora clone | 97 |
| 21 | KY979289.1 | Uncultured Glomus clone | 99 |
| 22 | KY232529.1 | Uncultured Glomus clone | 99 |
| 23 | HE613489.1 | Uncultured Paraglomus clone | 99 |
| 24 | KU668988.1 | Uncultured Claroideoglomu | 99 |
表5 2017年土壤AM真菌DGGE条带对比结果 |
| 条带 编号 | GeneBank 登录号 | 同源性最高的菌株 | 相似度/% |
|---|---|---|---|
| 1 | KY979378.1 | Uncultured Glomus clone | 100 |
| 2 | KY232438.1 | Uncultured Glomus clone | 99 |
| 3 | KY979384.1 | Uncultured Glomus clone | 100 |
| 4 | KX154256.1 | Uncultured Rhizophagus clone | 100 |
| 5 | KY173792.1 | Uncultured Glomus clone | 97 |
| 6 | KY232420.1 | Uncultured Glomus clone | 98 |
| 7 | LN621194.1 | Uncultured Claroideoglomus | 100 |
| 8 | MG835539.1 | Uncultured Rhizophagus clone | 100 |
| 9 | KU361755.1 | Uncultured Glomus clone | 99 |
| 10 | KU668988.1 | Uncultured Claroideoglomus | 100 |
| 11 | KY232471.1 | Uncultured Glomus clone | 100 |
| 12 | MF567532.1 | Uncultured Glomeromycotina clone | 99 |
| 13 | HG425740.1 | Uncultured Glomus clone | 100 |
| 14 | KX462871.1 | Uncultured Rhizophagus clone | 99 |
| 15 | KF601851.1 | Uncultured Glomus clone | 100 |
| 16 | KT238942.1 | Uncultured Glomeromycotina clone | 100 |
| 17 | KY979306.1 | Uncultured Glomus clone | 100 |
| 18 | KY232617.1 | Uncultured Glomus clone | 99 |
| 19 | MF567532.1 | Uncultured Glomeromycotina clone | 100 |
| 20 | LT672514.1 | Uncultured Claroideoglomus | 99 |
| 21 | KY232529.1 | Uncultured Glomus clone | 100 |
表6 土壤AM真菌DGGE图谱多样性指数 |
| 种植年份 | 种植地点 | 品种 | 香农-威纳指数(H') | 丰富度(S) | 均匀度指数(J) |
|---|---|---|---|---|---|
| 2016年 | 河北省 | DBN9936 | 2.94±0.12Aa | 21.67±1.53Aa | 0.96±0.02Aa |
| DBN318 | 2.94±0.12Aa | 21.67±1.53Aa | 0.96±0.02Aa | ||
| 吉林省 | DBN9936 | 2.83±0.14Aa | 20.67±2.08Aa | 0.94±0.02Aa | |
| DBN318 | 2.83±0.14Aa | 20.67±2.08Aa | 0.94±0.02Aa | ||
| 2017年 | 河北省 | DBN9936 | 3.23±0.02Ab | 29.33±0.58Ab | 0.95±0.01Aa |
| DBN318 | 3.10±0.11Ab | 26.33±2.31Ab | 0.95±0.01Aa | ||
| 吉林省 | DBN9936 | 3.33±0.10Aa | 31.67±2.08Aa | 0.96±0.10Aa | |
| DBN318 | 3.28±0.11Aa | 30.33±2.31Aa | 0.96±0.01Aa |
| 注：同一列数据下的英文大写字母表示同一年份同一地点下不同品种间的多重比较，小写字母表示同一品种同一年份下不同地点间的多重比较。字母不同表示处理间某指数差异达显著水平(P<0.05)。下同。 |
表7 转基因玉米对土壤微生物PLFA含量的影响 |
| 种植地点 | 品种 | 2016年总PLFA含量/(nmol/g) | 2017年总PLFA含量/(nmol/g) |
|---|---|---|---|
| 河北省 | DBN9936 | 45.19±2.93Ab | 45.27±3.02Ab |
| DBN318 | 48.13±3.46Ab | 44.13±4.01Ab | |
| 吉林省 | DBN9936 | 55.59±3.44Aa | 63.16±6.42Aa |
| DBN318 | 59.42±3.65Aa | 55.14±3.75Aa |
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A 52-week feeding study in cynomolgus macaques was carried out to evaluate the safety of Bt rice Huahui 1 (HH1), a transgenic rice line expressing Cry1Ab/1Ac protein. Monkeys were fed a diet with 20% or 60% HH1 rice, 20% or 60% parental rice (Minghui 63, MH63), normal diet, normal diet spiked with purified recombinant Cry1Ab/1Ac fusion protein or bovine serum albumin (BSA) respectively. During the feeding trail, clinical observations were conducted daily, and multiple parameters, including body weight, body temperature, electrocardiogram, hematology, blood biochemistry, serum metabolome and gut microbiome were examined at regular intervals. Upon sacrifice, the organs were weighted, and the macroscopic, microscopic and electron microscopic examinations were performed. The results show no adverse or toxic effects of Bt rice HH1 or Cry1Ab/1Ac fusion protein on monkeys. Therefore, the present 52-week primate feeding study suggests that the transgenic rice containing Cry 1Ab/1Ac is equivalent to its parental rice line MH63. Copyright © 2016 Elsevier Ltd. All rights reserved.
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Large quantities of Bacillus thuringiensis (Bt) corn plant residue are left in the field after harvest, which may have implications for the soil ecosystem. Potential impacts on soil organisms will also depend on the persistence of the Bt toxin in plant residues. Therefore, it is important to know how long the toxin persists in plant residues. In two field studies in the temperate corn-growing region of Switzerland we investigated degradation of the Cry1Ab toxin in transgenic Bt corn leaves during autumn, winter and spring using an enzyme-linked immunosorbent assay (ELISA). In the first field trial, representing a tillage system, no degradation of the Cry1Ab toxin was observed during the first month. During the second month, Cry1Ab toxin concentrations decreased to approximately 20% of their initial values. During winter, there was no further degradation. When temperatures again increased in spring, the toxin continued to degrade slowly, but could still be detected in June. In the second field trial, representing a no-tillage system, Cry1Ab toxin concentrations decreased without initial delay as for soil-incorporated Bt plants, to 38% of the initial concentration during the first 40 days. They then continued to decrease until the end of the trial after 200 days in June, when 0.3% of the initial amount of Cry1Ab toxin was detected. Our results suggest that extended pre- and post-commercial monitoring are necessary to assess the long-term impact of Bt toxin in transgenic plant residues on soil organisms.
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Root systems of most land plants form arbuscular mycorrhizal (AM) symbioses in the field, and these contribute to nutrient uptake. AM roots have two pathways for nutrient absorption, directly through the root epidermis and root hairs and via AM fungal hyphae into root cortical cells, where arbuscules or hyphal coils provide symbiotic interfaces. New physiological and molecular evidence shows that for phosphorus the mycorrhizal pathway (MP) is operational regardless of plant growth responses (positive or negative). Amounts delivered cannot be determined from plant nutrient contents because when responses are negative the contribution of the direct pathway (DP) is reduced. Nitrogen (N) is also delivered to roots via an MP, but the contribution to total N requirement and the costs to the plant are not clear. The functional interplay between activities of the DP and MP has important implications for consideration of AM symbioses in ecological, agronomic, and evolutionary contexts.
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梁晋刚, 焦悦, 刘鹏程, 等. 丛枝菌根真菌作为指示性物种评估转基因作物对土壤微生物影响的研究概述[J]. 浙江农业学报, 2018, 30(7): 1267-1272.
转基因作物与土壤生态系统环境要素紧密相关。土壤微生物作为土壤生态系统的一个重要组成部分,转基因作物大面积种植是否会给土壤微生物带来影响已成为其生态风险评价中不可忽视的一个方面。土壤中的微生物种类繁多,因此在研究时有必要选取一些具有代表性的微生物进行分析。丛枝菌根真菌作为一类重要的土壤环境指示微生物,在改善土壤质量与健康状况、提高生物多样性、促进植物生长等方面发挥着不可替代的作用。文章就主要转基因作物对丛枝菌根真菌的影响和作用机制进行综述,认为可将丛枝菌根真菌作为评价转基因作物对土壤生态系统影响的指示生物。
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The commercial use of genetically modified (GM) plants has significantly increased worldwide. The interactions between GM plants and arbuscular mycorrhizal (AM) fungi are of considerable importance given the agricultural and ecological role of AM and the lack of knowledge regarding potential effects of drought-tolerant GM corn ( L.) on AM fungal symbiosis. This work studied AM fungal colonization in five corn lines growing under two different irrigation regimes (30 and 100% of soil field capacity [SFC]). Four of the lines were GM corn, and two of these were drought tolerant. The experiment was conducted for 60 d in a growth chamber under constant irrigation, after which mycorrhization, corn biomass, and days to plant senescence (DTS) were evaluated. Arbuscular mycorrhizal fungal species of the order were predominant in the soil inocula. At the end of the experiment, all plants showed AM colonization. Mycorrhization was higher at 30% SFC than at 100% SFC. Within the same corn line, the AM fungi produced more vesicles in plant roots under drought stress. Among treatments, DTS varied significantly, and drought-tolerant GM corn lines survived longer than the wild-type corn when maintained at 100% SFC. Corn biomass did not vary among treatments, and no correlations were found between DTS or biomass and mycorrhization. We conclude that overexpression of the gene in corn plants under the experimental conditions of this study did not affect AM fungal infectivity and improved the tolerance of the corn to drought stress.Copyright © by the American Society of Agronomy, Crop Science Society of America, and Soil Science Society of America, Inc.
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郭静, 罗培宇, 杨劲峰, 等. 长期施肥对棕壤丛枝菌根真菌群落结构及其侵染的影响[J]. 中国农业科学, 2018, 51(24):4677-4689.
目的 丛枝菌根(Arbuscular mycorrhizal, AM)真菌有改善根际土壤环境、促进植物对养分的吸收、增强植物抗逆性和增加农作物产量等重要作用。本研究旨在探明长期施肥条件下玉米-大豆轮作棕壤丛枝菌根真菌群落结构、对玉米根系侵染的变化及其影响因素。方法 以沈阳农业大学棕壤肥料长期定位试验（38年）耕层（0—20 cm）土壤为材料,于2016年6月选取其中6个施肥处理：（1）不施肥处理（CK）;（2）单施化学氮肥（N）;（3）施用化学氮磷肥（NP）;（4）施用化学氮磷钾肥（NPK）;（5）单施有机肥（M）;（6）有机肥和化学氮磷肥配施（MNP）,采用PCR-DGGE、克隆测序及台盼蓝染色法,分析土壤和玉米根系定殖的AM真菌群落结构及侵染率,并结合环境因素进行冗余分析和典型对应分析。结果 施用有机肥处理土壤的碱解氮（AHN）、速效磷（AP）、速效钾（AK）、铵态氮（NH4 +-N）、硝态氮（NO3 --N）和可溶性有机碳（DOC）含量显著高于单施化肥和不施肥处理,且趋势为：有机肥处理>化肥处理>不施肥处理;与不施肥处理相比,单施化肥处理显著降低了土壤pH值,而施用有机肥处理显著提高了土壤pH值。通过PCR-DGGE及割胶测序,从土壤中得到AM真菌条带22条,根系AM真菌条带仅9条,共分离出13个OTU,从土壤样品中分离的AM真菌种群主要为球囊霉菌和巨孢囊霉属,而侵染玉米根系的AM真菌只有球囊霉菌。聚类分析表明长期不同施肥将棕壤中AM真菌分为了三大类群,分别为单施氮肥处理、施用有机肥处理和其他处理;根系AM真菌分为三大类群,第一类群NPK处理、第二类群为M处理和NP处理、第三类群为其他施肥处理。施用有机肥处理AM真菌的孢子密度显著高于单施化肥和不施肥处理,趋势为：有机肥处理﹥化肥处理﹥不施肥处理。各施肥处理AM真菌侵染率趋势为：NPK处理>施用有机肥处理>其他施肥处理。冗余分析结果表明棕壤AM真菌多样性与土壤理化性质无相关性,而孢子密度与土壤AHN、NH4 +-N、AP、AK、DOC及土壤含水量呈显著正相关;侵染率与土壤NO3 --N呈显著正相关;侵染率与孢子密度之间呈显著正相关;AM真菌的多样性与孢子密度和侵染率之间没有相关性。典型对应分析表明AHN、AK、DOC、 NH4 +-N 对AM真菌的群落组成影响显著。 结论 长期施肥通过改变土壤理化性质,从而对棕壤AM真菌的群落结构产生了显著影响,进而对AM真菌的侵染产生影响。
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Low availability of nitrogen (N) is often a major limiting factor to crop yield in most nutrient-poor soils. Arbuscular mycorrhizal (AM) fungi are beneficial symbionts of most land plants that enhance plant nutrient uptake, particularly of phosphate. A growing number of reports point to the substantially increased N accumulation in many mycorrhizal plants; however, the contribution of AM symbiosis to plant N nutrition and the mechanisms underlying the AM-mediated N acquisition are still in the early stages of being understood. Here, we report that inoculation with AM fungus remarkably promoted rice () growth and N acquisition, and about 42% of the overall N acquired by rice roots could be delivered via the symbiotic route under N-NO supply condition. Mycorrhizal colonization strongly induced expression of the putative nitrate transporter gene in rice roots, and its orthologs in and in OsNPF4.5 is exclusively expressed in the cells containing arbuscules and displayed a low-affinity NO transport activity when expressed in oocytes. Moreover, knockout of resulted in a 45% decrease in symbiotic N uptake and a significant reduction in arbuscule incidence when NO was supplied as an N source. Based on our results, we propose that the NPF4.5 plays a key role in mycorrhizal NO acquisition, a symbiotic N uptake route that might be highly conserved in gramineous species.Copyright © 2020 the Author(s). Published by PNAS.
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Soilborne pathogens can contribute to diversity maintenance in tree communities through the Janzen-Connell effect, whereby the pathogenic reduction of seedling performance attenuates with distance from conspecifics. By contrast, arbuscular mycorrhizal fungi (AMF) have been reported to promote seedling performance; however, it is unknown whether this is also distance dependent. Here, we investigate the distance dependence of seedling performance in the presence of both pathogens and AMF. In a subtropical forest in south China, we conducted a four-year field census of four species with relatively large phylogenetic distances and found no distance-dependent mortality for newly germinated seedlings. By experimentally separating the effects of AMF and pathogens on seedling performance of six subtropical tree species in a shade house, we found that soil pathogens significantly inhibited seedling survival and growth while AMF largely promoted seedling growth, and these effects were host specific and declined with increasing conspecific distance. Together, our field and experimental results suggest that AMF can neutralize the negative effect of pathogens and that the Janzen-Connell effect may play a less prominent role in explaining diversity of nondominant tree species than previously thought.
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Contents Summary 1059 I. Introduction: pathways of influence and pervasiveness of effects 1060 II. AM fungal richness effects on ecosystem functions 1062 III. Other dimensions of biodiversity 1062 IV. Back to basics - primary axes of niche differentiation by AM fungi 1066 V. Functional diversity of AM fungi - a role for biological stoichiometry? 1067 VI. Past, novel and future ecosystems 1068 VII. Opportunities and the way forward 1071 Acknowledgements 1072 References 1072 SUMMARY: Arbuscular mycorrhizal (AM) fungi play important functional roles in ecosystems, including the uptake and transfer of nutrients, modification of the physical soil environment and alteration of plant interactions with other biota. Several studies have demonstrated the potential for variation in AM fungal diversity to also affect ecosystem functioning, mainly via effects on primary productivity. Diversity in these studies is usually characterized in terms of the number of species, unique evolutionary lineages or complementary mycorrhizal traits, as well as the ability of plants to discriminate among AM fungi in space and time. However, the emergent outcomes of these relationships are usually indirect, and thus context dependent, and difficult to predict with certainty. Here, we advocate a fungal-centric view of AM fungal biodiversity-ecosystem function relationships that focuses on the direct and specific links between AM fungal fitness and consequences for their roles in ecosystems, especially highlighting functional diversity in hyphal resource economics. We conclude by arguing that an understanding of AM fungal functional diversity is fundamental to determine whether AM fungi have a role in the exploitation of marginal/novel environments (whether past, present or future) and highlight avenues for future research.© 2018 The Authors. New Phytologist © 2018 New Phytologist Trust.
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The cultivation of genetically modified (GM) crops has increased significantly over the last decades. However, concerns have been raised that some GM traits may negatively affect beneficial soil biota, such as arbuscular mycorrhizal fungi (AMF), potentially leading to alterations in soil functioning. Here, we test two maize varieties expressing the Bacillus thuringiensis Cry1Ab endotoxin (Bt maize) for their effects on soil AM fungal communities. We target both fungal DNA and RNA, which is new for AM fungi, and we use two strategies as an inclusive and robust way of detecting community differences: (i) 454 pyrosequencing using general fungal rRNA gene-directed primers and (ii) terminal restriction fragment length polymorphism (T-RFLP) profiling using AM fungus-specific markers. Potential GM-induced effects were compared to the normal natural variation of AM fungal communities across 15 different agricultural fields. AM fungi were found to be abundant in the experiment, accounting for 8% and 21% of total recovered DNA- and RNA-derived fungal sequences, respectively, after 104 days of plant growth. RNA- and DNA-based sequence analyses yielded most of the same AM fungal lineages. Our research yielded three major conclusions. First, no consistent differences were detected between AM fungal communities associated with GM plants and non-GM plants. Second, temporal variation in AMF community composition (between two measured time points) was bigger than GM trait-induced variation. Third, natural variation of AMF communities across 15 agricultural fields in The Netherlands, as well as within-field temporal variation, was much higher than GM-induced variation. In conclusion, we found no indication that Bt maize cultivation poses a risk for AMF.
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崔红娟, 束长龙, 宋福平, 等. 转cry1ah基因玉米对根际土壤微生物群落结构的影响[J]. 东北农业大学学报, 2011, 42(7):30-38.
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文章所在专题
表1 试验地土壤基本理化性质表2 PCR反应的引物及反应条件表3 磷脂脂肪酸类群分类
图1 不同种植地点和年份下转基因和非转基因玉米土壤AM真菌DGGE图谱表4 2016年土壤AM真菌DGGE条带对比结果表5 2017年土壤AM真菌DGGE条带对比结果图2 土壤AM真菌群落结构相似性UPGMA聚类分析图3 土壤AM真菌系统发育树（2016年）图4 土壤AM真菌系统发育树（2017年）表6 土壤AM真菌DGGE图谱多样性指数表7 转基因玉米对土壤微生物PLFA含量的影响图5 土壤微生物类群PLFA含量的相对丰度/
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