复合污染尾矿废水中真菌群落多样性及其驱动机制

doi:10.17520/biods.2020181

摘要/Abstract

摘要：

金属尾矿废水中含有重金属以及多种有机和无机污染物, 然而在该极端生境中仍然有大量微生物存在。为了揭示碱性尾矿废水中真菌群落的组成模式和多样性格局及其维持机制, 本文利用ITS1区rDNA基因扩增子测序和qPCR技术对山西中条山十八河尾矿库废水中5个不同采样点真菌群落的组成、丰度和分布格局进行了研究。通过主坐标分析(PCoA)比较不同采样点间群落结构的差异性; 通过冗余分析(RDA)探讨了水体理化因子对真菌群落结构的影响; 通过零模型分析了影响群落结构的主要因素; 通过网络图分析了真菌类群之间的种间相互作用。结果表明, 布勒掷孢酵母属(Bullera)、Schizangiella、支顶孢属(Acremonium)和亚罗酵母属(Yarrowia)是主要的优势属, 真菌群落在不同采样地点从门到属水平的相对丰度均有明显变化。真菌群落丰度沿水流方向逐渐增加且与有机碳(TOC)浓度呈显著正相关。真菌群落的α-多样性与pH、重金属(As和Cu)、无机碳(IC)和铵态氮(NH₄⁺)浓度显著相关。真菌群落的空间结构在不同采样点具有明显差异, 这种差异性与理化因子没有显著关系; 不同采样点真菌群落的零偏差值均大于零, 且不同物种之间存在复杂的种间相互作用。以上结果说明, 在尾矿废水中环境因子只对真菌群落的α-多样性有显著影响, 而群落的β-多样性主要受种间相互作用关系的影响, 表明在碱性铜尾矿废水中存在比较复杂的真菌群落动态模式。

关键词: 尾矿废水, 真菌群落, 多样性, 维持机制

Abstract

Aims:Metal tailings drainage contains heavy metals and a variety of organic and inorganic pollutants, but a large number of microorganisms still exist in this extreme habitat. In order to reveal the composition pattern and diversity maintaining mechanism of fungal communities in alkaline copper mine drainage, the composition, abundance and distribution pattern of fungal communities in five water sites in Shibahe tailings reservoir were studied by using the rDNA gene amplification sequencing and qPCR in ITS1 area.
Methods:The fungal community compositions among different sample points were compared by principal coordinate analysis (PCoA). The relationship between physicochemical factors and the distribution of fungal communities were analyzed by redundancy analysis (RDA). The main factors influencing community structure were analyzed by null model. The interactions between fungal genera were analyzed by network.
Results:the results showed that Bullera, Schizangiella, Acremonium and Yarrowia were the dominant genera. The relative abundance of fungal communities varied significantly from phylum to genus level in different sampling sites. The abundance of fungal communities increased gradually along the direction of water flow and was positively correlated with the concentration of total organic carbon (TOC). PH, heavy metals (As and Cu), inorganic carbon (IC) and ammonium nitrogen (NH₄⁺) were significantly correlated with the α-diversity of the communities. The distribution of fungal communities in different sample sites was obviously different. The physicochemical factors had no significant effect on the distribution of fungal community. The null deviation values of fungal communities in different sampling sites were greater than zero, and there were complex interactions among different genera.
Conclusions:These results indicated that the environmental factors only have a significant correlation with the α-diversity of the fungal communities, while the β-diversity was mainly affected by the interaction between species in tailings drainage. Our findings highlight that there is a relatively complex fungal community dynamic pattern in alkaline copper tailings drainage.

Key words: tailings drainage, fungal communities, diversity, maintenance mechanism

刘晋仙, 柴宝峰, 罗正明 (2021) 复合污染尾矿废水中真菌群落多样性及其驱动机制. 生物多样性, 29, 373-384. DOI: 10.17520/biods.2020181.

Jinxian Liu, Baofeng Chai, Zhengming Luo (2021) Driving forces and the diversity of fungal communities in complex contaminated tailings drainage. Biodiversity Science, 29, 373-384. DOI: 10.17520/biods.2020181.

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链接本文: https://www.biodiversity-science.net/CN/10.17520/biods.2020181

https://www.biodiversity-science.net/CN/Y2021/V29/I3/373

图/表 9

表1 不同采样点水体理化参数(平均值±标准误) (n = 3)

Table 1 Water physical and chemical parameters in different sample sites (mean±SE) (n = 3)

因子 Parameter	采样点 Sample site
	STW1	STW2	STW3	SUSW	SDSW
pH	9.382 ± 0.095 ^a	9.131 ± 0.053 ^a	8.147 ± 0.048 ^b	8.190 ± 0.032 ^b	8.014 ± 0.076 ^b
溶解氧 DO (mg/L)	8.560 ± 0.195 ^a	10.218 ± 0.466 ^a	10.661 ± 0.527 ^a	10.643 ± 0.281 ^a	11.114 ± 3.292 ^a
电导率 EC (uS/cm)	1,834.333 ± 31.205 ^a	1,832.001 ± 20.466 ^a	865.143 ± 3.106 ^b	1,427.333 ± 59.218 ^ab	404.333 ± 8.511 ^c
硝态氮$NO_{3}^{-}$ (mg/L)	134.373 ± 5.417 ^a	85.170 ± 1.553 ^ab	14.315 ± 1.142 ^bc	5.265 ± 0.345 ^c	4.698 ± 0.364 ^c
亚硝态氮$NO_{2}^{-}$ (mg/L)	10.548 ± 0.405 ^a	6.959 ± 0.20 ^ab	1.259 ± 0.096 ^bc	0.575 ± 0.057 ^c	0.478 ± 0.044 ^c
铵态氮$NH_{4}^{+}$ (mg/L)	1.453 ± 0.003 ^ab	1.715± 0.019^a	0.340 ± 0.116 ^b	2.333 ± 0.030 ^a	0.387 ± 0.015 ^b
总碳 TC (mg/L)	20.600 ± 0.035 ^c	24.723 ± 0.818 ^bc	34.523 ± 3.215 ^ab	25.070 ± 0.023 ^bc	52.315 ± 0.136 ^a
总有机碳 TOC (mg/L)	7.597 ± 0.003 ^abc	8.520 ± 1.529 ^bc	14.100 ± 0.075 ^ab	5.253 ± 0.020 ^c	19.344 ± 3.374 ^a
无机碳 IC (mg/L)	13.005 ± 0.032 ^c	16.202 ± 0.291 ^bc	20.179 ± 2.703 ^bc	19.820 ± 0.001 ^ab	43.015 ± 0.061 ^a
硫酸盐$SO_{4}^{2-}$ (mg/L)	1,582.500 ± 2.977 ^a	1,207.558 ± 100.997 ^a	837.255 ± 38.754 ^ab	897.900 ± 97.900 ^ab	117.655 ± 11.311 ^c
砷 As (mg/L)	2.407± 0.021^a	2.853 ± 0.171 ^a	0.155 ± 0.012 ^b	0.180 ± 0.009 ^ab	0.003 ± 0.003 ^b
镉 Cd (mg/L)	0.004 ± 0.001 ^a	0.002 ± 0.001 ^a	0.002 ± 0.003 ^a	0.003 ± 0.002 ^a	BDL
铜 Cu (mg/L)	0.017 ± 0.009 ^ab	0.032 ± 0.004 ^a	0.016 ± 0.004 ^ab	0.006 ± 0.002 ^b	0.008 ± 0.001 ^b
铅 Pb (mg/L)	0.060 ± 0.036 ^a	0.021 ± 0.013 ^a	0.025 ± 0.030 ^a	0.043 ± 0.006 ^a	BDL
锌 Zn (mg/L)	0.012 ± 0.001 ^ab	0.011 ± 0.004 ^b	0.009 ± 0.003 ^b	2.741 ± 1.058 ^a	0.249 ± 0.057 ^a

表1 不同采样点水体理化参数(平均值±标准误) (n = 3)

Table 1 Water physical and chemical parameters in different sample sites (mean±SE) (n = 3)

因子 Parameter	采样点 Sample site
	STW1	STW2	STW3	SUSW	SDSW
pH	9.382 ± 0.095 ^a	9.131 ± 0.053 ^a	8.147 ± 0.048 ^b	8.190 ± 0.032 ^b	8.014 ± 0.076 ^b
溶解氧 DO (mg/L)	8.560 ± 0.195 ^a	10.218 ± 0.466 ^a	10.661 ± 0.527 ^a	10.643 ± 0.281 ^a	11.114 ± 3.292 ^a
电导率 EC (uS/cm)	1,834.333 ± 31.205 ^a	1,832.001 ± 20.466 ^a	865.143 ± 3.106 ^b	1,427.333 ± 59.218 ^ab	404.333 ± 8.511 ^c
硝态氮$NO_{3}^{-}$ (mg/L)	134.373 ± 5.417 ^a	85.170 ± 1.553 ^ab	14.315 ± 1.142 ^bc	5.265 ± 0.345 ^c	4.698 ± 0.364 ^c
亚硝态氮$NO_{2}^{-}$ (mg/L)	10.548 ± 0.405 ^a	6.959 ± 0.20 ^ab	1.259 ± 0.096 ^bc	0.575 ± 0.057 ^c	0.478 ± 0.044 ^c
铵态氮$NH_{4}^{+}$ (mg/L)	1.453 ± 0.003 ^ab	1.715± 0.019^a	0.340 ± 0.116 ^b	2.333 ± 0.030 ^a	0.387 ± 0.015 ^b
总碳 TC (mg/L)	20.600 ± 0.035 ^c	24.723 ± 0.818 ^bc	34.523 ± 3.215 ^ab	25.070 ± 0.023 ^bc	52.315 ± 0.136 ^a
总有机碳 TOC (mg/L)	7.597 ± 0.003 ^abc	8.520 ± 1.529 ^bc	14.100 ± 0.075 ^ab	5.253 ± 0.020 ^c	19.344 ± 3.374 ^a
无机碳 IC (mg/L)	13.005 ± 0.032 ^c	16.202 ± 0.291 ^bc	20.179 ± 2.703 ^bc	19.820 ± 0.001 ^ab	43.015 ± 0.061 ^a
硫酸盐$SO_{4}^{2-}$ (mg/L)	1,582.500 ± 2.977 ^a	1,207.558 ± 100.997 ^a	837.255 ± 38.754 ^ab	897.900 ± 97.900 ^ab	117.655 ± 11.311 ^c
砷 As (mg/L)	2.407± 0.021^a	2.853 ± 0.171 ^a	0.155 ± 0.012 ^b	0.180 ± 0.009 ^ab	0.003 ± 0.003 ^b
镉 Cd (mg/L)	0.004 ± 0.001 ^a	0.002 ± 0.001 ^a	0.002 ± 0.003 ^a	0.003 ± 0.002 ^a	BDL
铜 Cu (mg/L)	0.017 ± 0.009 ^ab	0.032 ± 0.004 ^a	0.016 ± 0.004 ^ab	0.006 ± 0.002 ^b	0.008 ± 0.001 ^b
铅 Pb (mg/L)	0.060 ± 0.036 ^a	0.021 ± 0.013 ^a	0.025 ± 0.030 ^a	0.043 ± 0.006 ^a	BDL
锌 Zn (mg/L)	0.012 ± 0.001 ^ab	0.011 ± 0.004 ^b	0.009 ± 0.003 ^b	2.741 ± 1.058 ^a	0.249 ± 0.057 ^a

表2 十八河尾矿废水中5个采样点真菌群落的丰富度和多样性(平均值±标准误) (n = 3)。

Table 2 Richness and diversity index of the fungal communities in five sample sites of tailings drainage from Shibahe (mean±SE) (n = 3)

采样点 Sampling point	序列数 Sequence number	OTU个数 OTUs	Chao1指数 Chao1 index	Shannon指数 Shannon index	Simpson指数 Simpson index	盖度 Coverage
STW1	76,565.673 ± 251.658 ^c	404.675 ± 13.436 ^c	478.636 ± 25.356 ^b	5.496 ± 0.012 ^a	0.948 ± 0.001 ^a	0.999 ± 0.000 ^a
STW2	63,438.606 ± 317.863 ^d	118.658 ± 6.813 ^e	135.338 ± 10.784 ^d	2.387 ± 0.019 ^e	0.554 ± 0.005 ^d	0.998 ± 0.001 ^a
STW3	80,413.333 ± 224.795 ^b	499.582 ± 12.000 ^a	519.275 ± 16.587 ^a	5.096 ± 0.015 ^b	0.938 ± 0.001 ^b	0.997 ± 0.001 ^b
SUSW	81,207.36±165.631^a	460.055 ± 6.084 ^b	484.696 ± 11.345 ^b	2.982 ± 0.017 ^c	0.586 ± 0.004 ^d	0.996 ± 0.001 ^b
SDSW	81,094.014±45.935^a	237.336 ± 19.759 ^d	259.418 ± 11.001 ^c	2.585 ± 0.013 ^d	0.692 ± 0.002 ^c	0.999 ± 0.001 ^a

图1 不同采样点真菌群落ITS rDNA拷贝数。STW1、STW2和STW3表示尾矿库中3个采样点, SUSW和SDSW表示尾矿库外2个渗流水采样点。

Fig. 1 Fungal ITS rDNA copy numbers at the different sample points. STW1, STW2 and STW3 represent three sample sites in the tailings reservoir, SUSW and SDSW represent two seepage water sample sites outside the tailings reservoir.

图2 不同采样点真菌群落在门水平的组成。STW1、STW2和STW3表示尾矿库中3个采样点, SUSW和SDSW表示尾矿库外2个渗流水采样点。

Fig. 2 Fungal community composition of each sampling point at the phyla level. STW1, STW2 and STW3 represent three samplingpoints in the tailings reservoir, SUSW and SDSW represent two seepage water sample sites outside the tailings reservoir.

图3 不同采样点真菌群落的优势类群。(a)纲水平; (b)目水平; (c)科水平; (d)属水平。STW1、STW2和STW3表示尾矿库中3个采样点, SUSW和SDSW表示尾矿库外2个渗流水采样点。

Fig. 3 Dominant fungi in different sample sites. (a) Class level; (b) Order level; (c) Family level; (d) Genus level. STW1, STW2 and STW3 represent three samplingpoints in the tailings reservoir, SUSW and SDSW represent two seepage water sample sites outside the tailings reservoir.

图4 真菌优势类群组成与环境因子的RDA分析。(a)门水平; (b)纲水平; (c)目水平; (d)科水平; (e)属水平。NH4+: 铵态氮; IC:无机碳; TOC: 总有机碳; Cu: 铜。

Fig. 4 RDA showing the relationship between environmental parameters and composition of dominant fungal taxa. (a) Phylum level; (b) Class level; (c) Order level; (d) Family level; (e) Genus level. $NH_{4}^{+}$: Ammonium nitrogen; IC: Inorganic carbon; TOC: Total organic carbon; Cu: Copper.

表3 真菌群落的α多样性指数与环境参数的相关性系数

Table 3 Correlation coefficient of α diversity index of fungal communities and environmental parameters

环境因子 Parameter	OTUs
pH	-0.706**	-0.710**	-0.258	-0.268	0.107
DO	-0.193	-0.179	-0.329	-0.404	0.057
EC	-0.338	-0.336	-0.007	-0.200	0.257
$NO_{3}^{-}$	-0.132	-0.129	0.346	0.293	0.493
$NO_{2}^{-}$	-0.132	-0.132	0.350	0.296	0.471
$NH_{4}^{+}$	-0.189	-0.198	-0.214	-0.536*	-0.307
TC	0.106	0.086	-0.343	-0.118	-0.004
TOC	0.060	0.080	-0.025	0.297	0.425*
IC	-0.203	-0.244	-0.530*	-0.434	-0.498
$SO_{4}^{2-}$	-0.262	-0.270	0.229	0.068	0.007
As	-0.628*	-0.632*	-0.186	-0.261	0.104
Cd	-0.376	-0.340	-0.004	0.082	-0.021
Cu	-0.616*	-0.579*	-0.400	-0.239	0.346
Pb	-0.126	-0.082	0.179	0.039	-0.190
Zn	0.179	0.161	-0.050	-0.146	-0.350

图5 真菌群落空间分布格局及其影响因素。(a)在OTU水平基于Bray-Curtis距离的真菌群落PCoA排序结果; (b)真菌群落结构与环境因子的冗余分析; (c)基于Bray-Curtis距离的零偏差结果。IC: 无机碳; STW1、STW2和STW3表示尾矿库中3个采样点, SUSW和SDSW表示尾矿库外2个渗流水采样点。

Fig. 5 Spatial distribution pattern of fungal community structure and its influencing factors. (a) PCoA analysis based on Bray-Curtis similarities of fungal communities at OTU level; (b) RDA showing the relationship between environmental parameters and fungal communities; (c) Null deviation result based on Bray-Curtis distance. IC: Inorganic carbon; STW1, STW2 and STW3 represent three sample sites in the tailings reservoir, SUSW and SDSW represent two seepage water sample sites outside the tailings reservoir.

图6 属水平真菌类群间相互作用网络图。每个节点代表一个属, M1?M6代表6个独立模块, 节点大小与丰度成正比, 有显著相关性的节点通过线连接起来, 红线表示正相关, 绿线表示负相关。

Fig. 6 Network interactions of fungal groups at genus level. Each circle represents a genus and M1-M6 represent the six independent modules in the network diagram. The sizes of the circles are proportional to the value of abundance. Lines connecting two nodes represent the interactions between them. Red lines represent the positive significant correlations and green lines represent a negative significant correlation.

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