Plant and arbuscular mycorrhizal fungal diversities in field margins of the globally important agricultural heritage rice-fish system
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摘要: 全球重要农业文化遗产系统(Globally Important Agricultural Heritage Systems, GIAHS)对农业生物多样性的保育作用近年来受到关注。研究已表明, GIAHS-青田稻鱼共生系统保持高的水稻品种多样性和田鱼遗传多样性, 但在保育野生植物资源及其相关生物方面的作用仍不清楚。本文研究了GIAHS-青田稻鱼共生系统农田边界栖息的植物及与之共生的丛枝菌根真菌(arbuscular mycorrhizal fungi, AMF)群落, 通过坡底、坡中和坡顶取样, 评估农田边界植物多样性, 并通过高通量测序测定农田边界土壤AMF群落的多样性和组成。结果表明, 在本研究所调查的范围内, 由于总高差不大, 坡底、坡中和坡顶3个样地的植物组成、AMF群落组成及土壤理化性质均无显著性差异。青田稻鱼共生系统的农田边界分布有53科109种维管植物, α多样性指数(Shannon-Wiener指数)为0.74~0.84,
β多样性指数(Whittaker指数)为0.67~0.77。农田边界植物AMF侵染率为6.67%~44.44%, 根际土壤AMF分属9科14属62种, 优势科为球囊霉科(Glomeraceae), 优势属为球囊霉属(Glomus); 农田边界的土壤碳、氮、磷、钾含量较高。相关分析表明, AMF群落多样性与植物多样性呈正相关, 与土壤碳、氮、磷、钾含量也呈正相关, 其中与土壤速效磷的相关性最显著。这些研究结果表明, GIAHS-青田稻鱼共生系统农田边界保育有较高植物多样性及与之相共生的AMF群落, 研究结果有助于加深人们对GIAHS在生物多样性保育方面的理解。 Abstract: The role of Globally Important Agricultural Heritage Systems (GIAHS) in agro-biodiversity conservation is recently concerned. Studies have shown that the GIAHS-Qingtian rice-fish system has maintained a high diversity of rice varieties and local common carp population. The wild plant species diversity and the associated biota maintained in the GIAHS-Qingtian rice-fish system, however, are less well known. In this study, we examined plants and the associated arbuscular mycorrhizal fungi (AMF) in the field margins of the GIAHS-Qingtian rice-fish system. At the pilot site of the rice-fish system in Qingtian County, southern Zhejiang Province (120°26′–121°41′E, 27°25′–28°57′N), we investigated wild plant species diversity and collected soil samples from the rhizosphere at the field margin along a hillside with terrace fields. The α and β diversity indices of wild plants were calculated. The AMF communities in the soil samples were determined using high-throughput sequencing, and their diversity and composition were analyzed. No significant difference was found in the sample plots at different position along the slope. In the GIAHS-Qingtian rice-fish system, 109 vascular plant species belonging to 53 families in the field margin were found. Plant species α diversity (Shannon-Wiener index) was 0.74–0.84, and the β diversity (Whittaker index) was 0.67–0.77. The average AMF colonization rate of the plant species in the field margin was 6.67%–44.44%. Nine families, 14 genera, and 62 AMF species were found in soil along the field margin. The dominant AMF family was Glomeraceae, with a relative abundance of 87.29%–89.64%, and the dominant genus was Glomus (87.29%–89.64%). Carbon, nitrogen, phosphorus, and potassium contents were relatively high in the rhizospheric soil at the field margin. Correlation analysis showed that the AMF species diversity index (Shannon index) was positively correlated with the plant species diversity index (species richness). At the genus level, the AMF species diversity index (Shannon index) of Glomus was positively correlated with total soil nitrogen, available nitrogen, organic carbon, and organic matter. At the operational taxonomic unit (OTU) level, available phosphorus content had the most significant effect on the soil AMF community among the soil properties tested based on redundancy analysis. Our results showed that high plant species diversity and associated AMF were conserved in the GIAHS-Qingtian rice-fish system. These results enhance our understanding of the important role of GIAHSs in maintaining the biodiversity of agricultural ecosystems. -
图 1 研究区域地理位置图(a: 浙江省行政区域, 陆上部分; b: 青田县区域; c: 龙现村内本次调查研究的区域)
Figure 1. Geographic location map of the study area (a: administrative region of Zhejiang Province, land part only; b: Qingtian County area; c: the study area in Longxian Village)
图 2 采样样地、样方分布图(a)和样方位置(b)
Figure 2. Diagram of sampling settings in Longxian Village, Qingtian, Zhejiang, China
图 4 农田边界土壤科(a)、属(b)分类水平上丛枝菌根真菌(AMF)群落组成图
Figure 4. Species composition diagram of arbuscular mycorrhizal fungi (AMF) community at the taxonomic level of family (a) and genus (b) in soil of field margin
图 5 丛枝菌根真菌群落物种组成在属水平(a)、OTU水平(b)上与土壤环境因子的Spearman相关系数热图
图中, X轴为环境因子。MC: 土壤含水量; TN: 土壤总氮含量; AN: 土壤碱解氮含量; OC: 土壤有机碳含量; OM: 土壤有机质含量; AP: 土壤有效磷含量; AK: 土壤速效钾含量。图的左侧和上侧为真菌物种及土壤环境因子的聚类树。图a和图b的右侧色块集内, 不同R值用不同的颜色表示, R值自深红(相关系数为1.0)至深蓝(相关系数为−1.0)变化, 以不同颜色展示不同R值。图中矩形色块中, *表示P≤0.05, **表示P≤0.01, ***表示P≤0.001。
Figure 5. Spearman correlation coefficient heat maps between species composition of arbuscular mycorrhizal fungi (AMF) community classified at genus level (a) and OTU level (b) and soil environmental factors
X axis represents various environmental factors. MC: moisture content; TN: total nitrogen; AN: alkali-hydrolyzed nitrogen; OC: organic carbon; OM: organic matter; AP: available phosphorus; AK: available potassium. R value is shown with different color rectangles in the figure. Clusters of AMF species and soil factors are on the left and top of Figures a and b. The right colored rectangles in the right part represents the relationship index of AMF species (a) and OTU (b) with soil factors varing from the dark red (R=1.0) to the dark blue (R=−1.0). If P≤0.05, the colored rectangle is marked with *, if P≤0.01, the rectangle is marked with **, and if P≤0.001, the rectangle is marked with ***.
表 1 不同坡位农田边界植物群落α多样性指数
Table 1. Species α diversity indexes of vegetation community in the field margins in different positions of hillside
位置
Location丰富度指数
Richness indexShannon-Wiener指数
Shannon-Wiener indexSimpson指数
Simpson indexPielou指数
Pielou index坡底 Base of hillside 20.33±3.03ns 0.84±0.07ns 0.77±0.05ns 0.67±0.06ns 坡中 Middle of hillside 18.17±1.22ns 0.78±0.05ns 0.73±0.02ns 0.62±0.03ns 坡顶 Top of hillside 15.00±2.08ns 0.74±0.14ns 0.72±0.07ns 0.63±0.08ns 数值为平均值±标准误差, n=3; ns表示不同坡位间差异未达显著水平。Value is mean±standard error, n=3; ns means no significant differences among the locations. 
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表 2 不同坡位农田边界植物群落β多样性指数
Table 2. Species β diversity of vegetation community in the field margins in different positions of hillside
位置 Location Jaccard指数 Jaccard index Whittaker指数 Whittaker index Sorenson指数 Sorenson index 坡底-坡中 Base-middle of hillside 0.13±0.03ns 0.77±0.05ns 0.23±0.05ns 坡中-坡顶 Middle-top of hillside 0.20±0.06ns 0.67±0.09ns 0.33±0.09ns 数值为平均值±标准误差, n=3; ns表示不同坡位间差异未达显著水平。Value is mean±standard error, n=3; ns means no significant difference among the locations.
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表 3 农田边界植物的丛枝菌根真菌(AMF)侵染率
Table 3. Arbuscular mycorrhizal fungi (AMF) colonization rates of plants in the field margin
宿主植物
Host plant根部总侵染率
Root colonization rate (%)络石 Trachelospermum jasminoides 44.44±0.04d 斑鸠菊 Vernonia esculenta 44.44±0.09d 地耳草 Hypericum japonicum 42.22±0.03d 堇菜 Viola verecunda 36.67±0.12cd 白茅 Imperata cylindrica 34.44±0.06cd 鸭跖草 Commelina communis 33.33±0.08bcd 野菊 Chrysanthemum indicum 31.11±0.07bcd 弓果黍 Cyrtococcum patens 28.89±0.11abcd 糯米团 Gonostegia hirta 26.67±0.08abcd 马唐 Digitaria sanguinalis 23.33±0.08abcd 狗尾草 Setaria viridis 22.22±0.02abcd 狗牙根 Cynodon dactylon 21.11±0.06abcd 败酱 Patrinia scabiosaefolia 17.78±0.07abc 酢浆草 Oxalis corniculata 10.00±0.04ab 积雪草 Centella asiatica 6.67±0.02a 数值为平均值±标准误差, n=3; 不同小写字母表示不同宿主植物间差异达显著水平(P<0.05)。Value is mean±standard error, n=3. Different lowercase letters mean significant differences among host plants.
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表 4 农田边界根际土壤碳氮磷钾特征
Table 4. Soil properties of field margin
位置
Location含水率Moisture
content (%)有机碳Organic carbon
(g∙kg−1)有机质Organic matter
(g∙kg−1)碱解氮Alkali-hydrolyzable nitrogen
(mg∙kg−1)有效磷Available phosphorus
(mg∙kg−1)速效钾Available potassium
(mg∙kg−1)全氮Total nitrogen
(g∙kg−1)坡底
Base of hillside45.12±3.04ab 33.75±4.31ns 58.18±7.42ns 276.36±25.8ns 26.34±5.93ns 176.05±12.13ns 4.36±0.42ns 坡中
Middle of hillside35.44±3.36a 25.36±1.63ns 43.72±2.81ns 205.16±14.35ns 58.29±28.37ns 181.58±8.02ns 2.55±0.19ns 坡顶
Top of hillside55.50±5.46b 33.13±1.76ns 57.12±3.03ns 260.47±11.23ns 41.59±2.13ns 161.54±5.65ns 3.14±0.09ns 数值为平均值±标准误差, n=3; ns表示不同坡位间差异未达显著水平。Value is mean±standard error, n=3; ns means no significant difference among the locations.
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表 5 土壤丛枝菌根真菌(AMF)群落丰富度与多样性指数(基于OTU)
Table 5. Species richness and diversity indexes of arbuscular mycorrhizal fungi (AMF) community in field margin (based on OTU)
位置
Location丰富度指数 Richness index 多样性指数 Diversity index 覆盖度
CoverageAce Chao Shannon Simpson 坡底 Base of hillside 73.01±7.34ns 72.35±7.96ns 2.99±0.17ns 0.09±0.02ns 0.99±0.01ns 坡中 Middle of hillside 68.06±6.54ns 68.10±6.68ns 2.85±0.13ns 0.11±0.02ns 0.99±0.01ns 坡顶 Top of hillside 59.87±3.92ns 58.39±3.86ns 2.49±0.40ns 0.17±0.08ns 0.99±0.01ns 数值为平均值±标准误差, n=3; ns表示不同坡位间差异未达显著水平。Value is mean±standard error, n=3; ns means no significant difference among the locations.
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表 6 植物物种丰富度和丛枝菌根真菌(AMF)群落Shannon指数相关性
Table 6. Correlation between plant species richness and Shannon index of arbuscular mycorrhizal fungi (AMF) community
相关系数 Correlation coefficient Pearson Spearman Kendall 物种丰富度 Species richness-Shannon index 0.703** 0.667** 0.518** **代表P<0.01 (双尾)水平相关性显著。** means signficant corrlation at P<0.01 level.
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