Ecology of Culturable Fungal Community in a GORE Cover Membrane System for Composting Agricultural Waste
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摘要:
目的 评估农业废弃物戈尔膜发酵过程中的真菌种群随时间和空间的变化规律，为农业废弃物的资源化利用和戈尔膜发酵效果评价提供理论依据。 方法 以农业废弃物戈尔膜发酵槽物料为研究对象，监测发酵过程物料的温度变化，并通过时间和空间格局采样，分离鉴定物料中的真菌种类及数量，分析发酵过程中的真菌种群数量分布、空间分布型指数、多样性指数和生态位特征等参数。 结果 发酵物料的温度监测表明，第2~16天的平均温度为55.27~74.64 ℃，定义为高温期；第17~27天的平均温度为41.26~50.64 ℃，定义为低温期。从72份物料样本中共分离鉴定出5种真菌，分别为沃尔夫被孢霉（Mortierella walfii）、青霉菌（Penicullium sp.）、烟曲霉（Aspergillus fumigatus）、棘曲霉（A. spinosus）和土曲霉（A. terreus）。从空间分布上看，烟曲霉在物料浅层和深层的种群数量均较多，分别为1.13×105 CFU·g−1和1.47×105 CFU·g−1；而棘曲霉在物料浅层和深层的种群数量均最少，分别为4.90×103 CFU·g−1和1.56×103 CFU·g−1，说明烟曲霉适宜在该物料中生长，而棘曲霉则不适宜。从发酵时间看，发酵前期 （3~12 d）的总菌量在4.10×104 ~1.30×105 CFU·g−1，发酵后期（17~27 d）的总菌量在9.35×103~2.63×104CFU·g−1。发酵过程中真菌种群呈现明显的差异性，且空间分布型为聚集分布，烟曲霉在物料中生存适应性最强，但竞争能力较弱或不存在竞争。 结论 农业废弃物戈尔膜发酵过程中的可培养真菌的数量在不同时间和空间上均存在明显的差异，其空间分布型为聚集分布。 Abstract:Objective Temporal and spatial characteristics of the culturable fungal community in a GORE cover membrane system (GCMS) for composting agricultural waste were analyzed to evaluate and enhance the utilization of the treatment system. Methods In a GCMS tank fermentation on agriculture waste material, temperature was continuously monitored, and varied temporal and spatial samples collected for microbial determination. Culturable fungi in the system were isolated, identified, and counted for analyzing and calculating spatial distribution pattern index, diversity index, and ecological niche characteristics. Results The average temperature of the composting material in the tank was 55.27-74.64 ℃ from the 2nd to the 16th day, which was defined as the high temperature period. In the low temperature period from the 17th to the 27th day, the average temperature was 41.26-50.64 ℃. From 72 specimens, 5 species of fungi, i.e., Mortierella walfii, Penicullium sp., Aspergillus fumigatus, Aspergillus spinosus, and Aspergillus terreus, were isolated and identified. Spatially, the population of A. fumigatus was the largest in both shallow and deep layers of the waste material with the counts of 1.13×105 CFU·g−1 and 1.47×105 CFU·g−1, respectively. Whereas A. aculeatus was the least in the same areas with the counts of 4.90×103 CFU·g−1 and 1.56×103 CFU·g−1, respectively. It indicated differentiated growth adaptabilities of the two fungal populations. Temporally, the total fungal counts were between 4.10×104 CFU·g−1 and 1.30×105 CFU·g−1 in the pre-fermentation period from the 3rd to the 12th day, and between 9.35×103 CFU·g−1 and 2.63×104CFU·g−1 in the post-fermentation period from the 17th to the 27th day. It appeared that the fungal community in the compost changed significantly during the fermentation process with a largely aggregated spatial distribution. A. fumigatus displayed the strongest growth adaptability under GCMS, even though it could be weak in competing with others, or simply because of a lack of competitive counterparts within the system. Conclusion The count of culturable fungi in GCMS differed significantly in different periods and locations. The spatial distribution of the fungal populations followed an aggregated pattern. -
Key words:
- GORE cover membrane system /
- fungi /
- spatial distribution /
- diversity index /
- niche characteristics
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图 1 农业废弃物戈尔膜发酵过程的温度变化
Figure 1. Temperature changes during fermentation of agricultural waste in GCMS
图 2 5个真菌种类的菌落形态
a：沃尔夫被孢霉；b：青霉菌； c：烟曲霉；d：棘曲霉；e：土曲霉。
Figure 2. Colony morphology of 5 fungal species
a: M. walfii; b: Penicullium sp.; c: A. fumigatus; d: A. spinosus; e: A. terreus.
表 1 农业废弃物戈尔膜发酵物料中的真菌分离与鉴定
Table 1. Isolation and identification of fungi from agricultural waste materials in GCMS
序号
No.真菌种类
Fungal species代表性菌株编号
Strain no.Genbank登录号
Genbank No.ITS BenA 1 沃尔夫被孢霉 Mortierella walfii FJAT-32629 OM831104 — 2 青霉菌 Penicullium sp. FJAT-32630 OM831105 OM897562 3 烟曲霉 Aspergillus fumigatus FJAT-32633 OM831106 OM897563 4 棘曲霉 Aspergillus spinosus FJAT-32654 OM831107 OM897564 5 土曲霉 Aspergillus terreus FJAT-32655 OM831108 OM897565 
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表 2 农业废弃物戈尔膜发酵过程不同样品的真菌种类与数量统计
Table 2. Fungal species and amount in samples from agricultural waste materials in GCMS
取样点
Sampling location真菌种类
Fungal species不同发酵时间的菌落数
Fungal amount at different fermentation times/（×102 CFU·g−1）0 d 3 d 6 d 9 d 12 d 17 d 21 d 24 d 27 d A浅层 A shallow layer 沃尔夫被孢霉 Mortierella walfii 30.0 10.0 0.0 0.0 0.0 2.0 7.3 1.5 5.6 青霉菌 Penicullium sp. 25.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 烟曲霉 Aspergillus fumigatus 4.0 0.0 163.0 41.0 33.5 59.4 87.5 7.0 0.0 棘曲霉 Aspergillus spinosus 7.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 土曲霉 Aspergillus terreus 0.0 0.0 0.0 5.5 0.0 10.0 0.0 0.0 0.0 A深层 A deep layer 沃尔夫被孢霉 Mortierella walfii 2.0 12.0 10.0 0.0 0.0 0.0 7.6 87.5 1.0 青霉菌 Penicullium sp. 10.0 2.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 烟曲霉 Aspergillus fumigatus 1.0 39.0 14.0 15.0 800.0 29.5 9.0 100.0 5.0 棘曲霉 Aspergillus spinosus 1.5 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 土曲霉 Aspergillus terreus 0.0 0.0 0.0 0.0 300.0 1.0 0.0 0.0 0.0 B浅层 B shallow layer 沃尔夫被孢霉 Mortierella walfii 0.0 10.0 1.0 0.0 0.0 0.0 0.5 0.6 0.6 青霉菌 Penicullium sp. 10.0 0.0 0.0 0.0 55.0 0.0 0.0 0.0 0.0 烟曲霉 Aspergillus fumigatus 0.0 16.0 270.0 0.0 0.0 0.0 1.0 0.4 0.7 棘曲霉 Aspergillus spinosus 20.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 土曲霉 Aspergillus terreus 0.0 0.0 0.0 180.0 0.0 0.0 0.0 0.0 0.0 B深层 B deep layer 沃尔夫被孢霉 Mortierella walfii 10.0 0.0 0.0 0.0 0.0 0.0 0.5 0.6 11.0 青霉菌 Penicullium sp. 0.0 0.0 0.0 0.0 0.0 0.0 0.4 0.0 0.0 烟曲霉 Aspergillus fumigatus 50.0 1.0 1.0 1.6 0.1 0.0 0.5 0.8 0.0 棘曲霉 Aspergillus spinosus 0.0 0.0 0.0 0.0 1.1 0.0 0.0 0.0 0.0 土曲霉 Aspergillus terreus 0.0 0.0 0.0 0.0 0.0 0.6 0.0 0.0 0.0 C浅层 C shallow layer 沃尔夫被孢霉 Mortierella walfii 3.0 4.0 1.0 0.0 0.0 0.0 0.5 8.8 11.0 青霉菌 Penicullium sp. 1.5 0.0 0.0 9.5 0.0 0.0 0.6 0.0 0.0 烟曲霉 Aspergillus fumigatus 9.5 23.0 1.5 0.0 11.0 0.0 1.3 0.0 2.0 棘曲霉 Aspergillus spinosus 0.0 1.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 土曲霉 Aspergillus terreus 0.0 0.0 0.0 420.0 0.0 1.0 0.0 0.0 0.0 C深层 C deep layer 沃尔夫被孢霉 Mortierella walfii 2.0 15.0 0.0 0.0 0.0 0.1 1.0 3.3 28.8 青霉菌 Penicullium sp. 8.0 0.0 0.0 0.0 0.0 0.0 0.6 0.0 0.0 烟曲霉 Aspergillus fumigatus 4.0 24.0 0.0 0.1 0.3 0.2 0.8 10.4 0.0 棘曲霉 Aspergillus spinosus 8.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 土曲霉 Aspergillus terreus 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 D浅层 D shallow layer 沃尔夫被孢霉 Mortierella walfii 5.0 35.0 1.0 0.0 0.0 1.0 9.3 20.0 0.0 青霉菌 Penicullium sp. 3.0 0.0 0.0 0.0 0.0 0.0 3.0 0.0 0.0 烟曲霉 Aspergillus fumigatus 1.0 170.0 24.0 135.0 29.0 3.2 25.5 6.6 0.0 棘曲霉 Aspergillus spinosus 11.0 10.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 土曲霉 Aspergillus terreus 0.0 0.0 0.0 0.0 0.0 1.0 0.0 0.0 0.0 D深层 D deep layer 沃尔夫被孢霉 Mortierella walfii 1.0 10.0 10.0 0.0 0.0 0.0 0.4 0.8 3.5 青霉菌 Penicullium sp. 25.0 0.0 0.0 0.0 0.0 0.0 1.8 0.0 0.0 烟曲霉 Aspergillus fumigatus 41.0 23.0 144.0 38.3 67.5 0.0 8.9 14.3 24.3 棘曲霉 Aspergillus spinosus 0.0 5.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 土曲霉 Aspergillus terreus 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0
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表 3 农业废弃物戈尔膜发酵过程真菌数量的空间分布
Table 3. Spatial distribution on fungal count in fermentation of agricultural waste in GCMS
真菌种类
Fungal species取样位置
Sampling location不同发酵时间的菌落数
Fungal amount at different fermentation times/（×102 CFU·g−1）0 d 3 d 6 d 9 d 12 d 17 d 21 d 24 d 27 d 总和 Total 烟曲霉 Aspergillus fumigatus 浅层 14.5 209.0 458.5 176.0 73.5 62.6 115.3 14.0 2.7 1126.1 深层 96.0 87.0 159.0 55.0 867.9 29.7 19.2 125.5 29.3 1468.6 棘曲霉 Aspergillus spinosus 浅层 38.0 11.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 49.0 深层 9.5 5.0 0.0 0.0 1.1 0.0 0.0 0.0 0.0 15.6 土曲霉 Aspergillus terreus 浅层 0.0 0.0 0.0 605.5 0.0 12.0 0.0 0.0 0.0 617.5 深层 0.0 0.0 0.0 0.0 300.0 1.6 0.0 0.0 0.0 301.6 沃尔夫被孢霉 Mortierella walfii 浅层 38.0 59.0 3.0 0.0 0.0 3.0 17.6 30.9 17.2 168.7 深层 15.0 37.0 20.0 0.0 0.0 0.1 9.5 92.2 44.3 218.1 青霉菌 Penicullium sp. 浅层 39.5 0.0 0.0 9.5 55.0 0.0 3.6 0.0 0.0 107.6 深层 43.0 2.0 0.0 0.0 0.0 0.0 2.8 0.0 0.0 47.8 总和 Total 293.5 410.0 640.5 846.0 1297.5 109.0 168.0 262.6 93.5 4120.6
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表 4 农业废弃物戈尔膜发酵前期真菌种群空间分布型指数
Table 4. Spatial distribution pattern index on fungal populations in agricultural waste in GCMS during pre-fermentation period
真菌种类
Fungal species取样位置
Sampling location拥挤度m*
Congestion degreeI指标
I indexm*/m指标
m*/m indexCA指标
CA index扩散系数C
Diffusion coefficientK指数
K index烟曲霉 Aspergillus fumigatus 浅层 34323.22 15693.22 1.84 0.84 15694.22 1.19 深层 72568.23 47270.23 2.87 1.87 47271.23 0.54 棘曲霉 Aspergillus spinosus 浅层 3746.35 2766.35 3.82 2.82 2767.35 0.35 深层 854.17 542.17 2.74 1.74 543.17 0.58 土曲霉 Aspergillus terreus 浅层 72659.00 60549.00 6.00 5.00 60550.00 0.20 深层 35999.00 29999.00 6.00 5.00 30000.00 0.20 沃尔夫被孢霉 Mortierella walfii 浅层 5666.50 3666.50 2.83 1.83 3667.50 0.55 深层 3100.81 1660.81 2.15 1.15 1661.81 0.87 青霉菌 Penicullium sp. 浅层 5098.59 3018.59 2.45 1.45 3019.59 0.69 深层 4921.22 4021.22 5.47 4.47 4022.22 0.22
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表 5 农业废弃物戈尔膜发酵后期真菌种群空间分布型指数
Table 5. Spatial distribution pattern index on fungal populations in agricultural waste in GCMS during post-fermentation period
真菌种类
Fungal species取样位置
Sampling location拥挤度m*
Congestion degreeI指标
I indexm*/m指标
m*/m indexCA指标
CA index扩散系数C
Diffusion coefficientK指数
K index烟曲霉 Aspergillus fumigatus 浅层 10310.28 5445.28 2.12 1.12 5446.28 0.89 深层 9991.55 4899.05 1.96 0.96 4900.05 1.04 土曲霉 Aspergillus terreus 浅层 2472.42 754.92 1.44 0.44 755.92 2.28 深层 8412.88 4760.38 2.30 1.30 4761.38 0.77 沃尔夫被孢霉 Mortierella walfii 浅层 449.00 359.00 4.99 3.99 360.00 0.25 深层 349.00 279.00 4.99 3.99 280.00 0.25 青霉菌 Penicullium sp. 浅层 10310.28 5445.28 2.12 1.12 5446.28 0.89 深层 9991.55 4899.05 1.96 0.96 4900.05 1.04
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表 6 农业废弃物戈尔膜发酵过程不同空间样本的真菌种群多样性指数
Table 6. Diversity index on fungal populations in spatial samples of agricultural waste in GCMS fermentation
空间样本
Spatial samples种类
Species数量
Amount/（×102 CFU·g−1）丰富度指数
Richness index (D)均匀度指数
Pielou’s evenness index(J')优势度指数
Simpson index (λ)0 d浅层 0 d shallow layer 4 130 0.32 0.96 0.72 3 d浅层 3 d shallow layer 3 279 0.20 0.61 0.39 6 d浅层 6 d shallow layer 2 461.5 0.09 0.06 0.01 9 d浅层 9 d shallow layer 3 791 0.18 0.54 0.36 12 d浅层 12 d shallow layer 2 128.5 0.11 0.98 0.49 17 d浅层 17 d shallow layer 3 77.6 0.22 0.53 0.32 21 d浅层 21 d shallow layer 3 136.5 0.21 0.46 0.27 24 d浅层 24 d shallow layer 2 44.9 0.12 0.90 0.43 27 d浅层 27 d shallow layer 2 19.9 0.13 0.57 0.23 0 d深层 0 d deep layer 4 163.5 0.31 0.76 0.57 3 d深层 3 d deep layer 4 131 0.32 0.59 0.48 6 d深层 6 d deep layer 2 179 0.10 0.51 0.20 9 d深层 9 d deep layer 1 55 0.00 0.00 0.00 12 d深层 12 d deep layer 3 1169 0.17 0.52 0.38 17 d深层 17 d deep layer 3 31.4 0.25 0.20 0.10 21 d深层 21 d deep layer 3 31.5 0.25 0.80 0.53 24 d深层 24 d deep layer 2 217.7 0.10 0.98 0.49 27 d深层 27 d deep layer 2 73.6 0.11 0.97 0.48
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表 7 农业废弃物戈尔膜发酵过程真菌种群生态位宽度和生态位重叠
Table 7. Niche breadth and overlap of fungal populations in spatial samples of agricultural waste in GCMS fermentation
种类
Species生态位宽度
Niche breadth生态位重叠Pianka指数
Niche overlap Pianka index烟曲霉
Aspergillus fumigatus棘曲霉
Aspergillus spinosus土曲霉
Aspergillus terreus沃尔夫被孢霉
Mortierella walfii青霉菌
Penicullium sp.烟曲霉 Aspergillus fumigatus 4.59 1.00 棘曲霉 Aspergillus spinosus 1.66 0.18 1.00 土曲霉 Aspergillus terreus 1.85 0.52 0.01 1.00 沃尔夫被孢霉 Mortierella walfii 4.64 0.33 0.45 0.00 1.00 青霉菌 Penicullium sp. 2.42 0.53 0.80 0.33 0.26 1.00
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