微生物学报

Sites	Longitude	Latitude	Altitude (m)	Overview of the surrounding environment of the sampling point
WA02	99°39′49″E	27°52′55″N	3 283.30	It’s a tourist area and a racecourse, with a large number of tourists
WA03	99°38′27″E	27°53′48″N	3 273.20	It’s a tourist area
WA04	99°38′9″E	27°54′8″N	3 284.60	It’s a mountainous area, and far from where people live
WA05	99°37′52″E	27°54′26″N	3 285.80	It’s a place where few people live, and there is foam in the water
WA06	99°37′43″E	27°54′18″N	3 285.69	It’s a planted forest, and far from where people live
WA07	99°37′48″E	27°53′50″N	3 274.55	It’s a natural pine forest, and far from where people live
WA08	99°37′43″E	27°53′36″N	3 272.30	It’s the mouth of the Naqu River, and far from where people live
WA09	99°37′26″E	27°53′53″N	3 272.30	It’s the cape of Napahai, with poor water mobility and mostly stagnant water, and there are people living nearby
WA10	99°37′17″E	27°53′8″N	3 272.30	It’s a spring hole in the water, with mountain spring water emerging, and far from the place where people live
WA11	99°37′52″E	27°51′43″N	3 272.30	There are many emergent plants in the water, and far from where people live
WA12	99°38′4″E	27°49′59″N	3 283.30	It’s a place with a large number of aquatic plants, and close to where people gather

Sites	Longitude	Latitude	Altitude (m)	Overview of the surrounding environment of the sampling point
WA02	99°39′49″E	27°52′55″N	3 283.30	It’s a tourist area and a racecourse, with a large number of tourists
WA03	99°38′27″E	27°53′48″N	3 273.20	It’s a tourist area
WA04	99°38′9″E	27°54′8″N	3 284.60	It’s a mountainous area, and far from where people live
WA05	99°37′52″E	27°54′26″N	3 285.80	It’s a place where few people live, and there is foam in the water
WA06	99°37′43″E	27°54′18″N	3 285.69	It’s a planted forest, and far from where people live
WA07	99°37′48″E	27°53′50″N	3 274.55	It’s a natural pine forest, and far from where people live
WA08	99°37′43″E	27°53′36″N	3 272.30	It’s the mouth of the Naqu River, and far from where people live
WA09	99°37′26″E	27°53′53″N	3 272.30	It’s the cape of Napahai, with poor water mobility and mostly stagnant water, and there are people living nearby
WA10	99°37′17″E	27°53′8″N	3 272.30	It’s a spring hole in the water, with mountain spring water emerging, and far from the place where people live
WA11	99°37′52″E	27°51′43″N	3 272.30	There are many emergent plants in the water, and far from where people live
WA12	99°38′4″E	27°49′59″N	3 283.30	It’s a place with a large number of aquatic plants, and close to where people gather

Sites	WA02	WA03	WA04	WA05	WA06	WA07	WA08	WA09	WA10	WA11	WA12
WA02	1
WA03	0.57	1
WA04	0.27	0.25	1
WA05	0.21	0.36	0.19	1
WA06	0.23	0.29	0.26	0.44	1
WA07	0.13	0.18	0.18	0.25	0.27	1
WA08	0.20	0.18	0.13	0.15	0.17	0.45	1
WA09	0.16	0.26	0.20	0.36	0.38	0.43	0.43	1
WA10	0.25	0.31	0.15	0.33	0.22	0.18	0.18	0.21	1
WA11	0.20	0.43	0.18	0.25	0.27	0.23	0.23	0.33	0.44	1
WA12	0.09	0.17	0.05	0.13	0.14	0.11	0.11	0.08	0.40	0.25	1

Sites	WA02	WA03	WA04	WA05	WA06	WA07	WA08	WA09	WA10	WA11	WA12
WA02	1
WA03	0.57	1
WA04	0.27	0.25	1
WA05	0.21	0.36	0.19	1
WA06	0.23	0.29	0.26	0.44	1
WA07	0.13	0.18	0.18	0.25	0.27	1
WA08	0.20	0.18	0.13	0.15	0.17	0.45	1
WA09	0.16	0.26	0.20	0.36	0.38	0.43	0.43	1
WA10	0.25	0.31	0.15	0.33	0.22	0.18	0.18	0.21	1
WA11	0.20	0.43	0.18	0.25	0.27	0.23	0.23	0.33	0.44	1
WA12	0.09	0.17	0.05	0.13	0.14	0.11	0.11	0.08	0.40	0.25	1

Sites	T/℃	Turb	pH	TP (mg/L)	TN (mg/L)	TOC (mg/L)	TH (mg/L)	EC (μS/cm)
The letters in the table indicate size and significant differences: “a” is the largest, “ab” is the second largest, and decreases in sequence from beginning to end; There is a significant difference between “a” “b” and “c”, with aP-value of < 0.05. “ab” indicates that there is no significant difference between “a” and “b”, while “bc” indicates that there is no significant difference between “b” and “c”, but there is a significant difference between “a” and “ab”, and so on.
WA02	11.0c	7.01bc	7.67c	0.06bc	0.99b	30.73c	189.00ab	343.00aab
WA03	15.0ab	6.51bc	7.90b	0.06bc	0.75bc	37.97bc	131.00b	257.00bc
WA04	16.0a	26.77ab	7.90b	0.09b	1.02ab	39.93ab	134.33ab	274.00b
WA05	16.0a	12.61ab	7.80bc	0.26a	2.06a	42.87ab	132.00ab	289.33ab
WA06	16.0a	34.07a	7.90b	0.14ab	1.29ab	40.63ab	136.00ab	283.00ab
WA07	16.0a	13.01ab	7.90b	0.11ab	0.97bc	41.07ab	124.33bc	279.33ab
WA08	16.0a	8.05b	8.00ab	0.07bc	0.71bc	38.07bc	127.00bc	277.00ab
WA09	16.0a	12.71ab	8.07ab	0.13ab	0.69bc	39.63bc	144.00ab	273.00bc
WA10	15.0ab	6.25bc	8.10a	0.10ab	0.56c	39.70b	117.00bc	267.33bc
WA11	15.0ab	4.30c	8.10a	0.06c	0.77bc	35.80bc	115.33c	251.00c
WA12	14.5b	21.90ab	7.90b	0.10ab	1.13ab	70.60a	228.00a	465.00a

Sites	T/℃	Turb	pH	TP (mg/L)	TN (mg/L)	TOC (mg/L)	TH (mg/L)	EC (μS/cm)
The letters in the table indicate size and significant differences: “a” is the largest, “ab” is the second largest, and decreases in sequence from beginning to end; There is a significant difference between “a” “b” and “c”, with aP-value of < 0.05. “ab” indicates that there is no significant difference between “a” and “b”, while “bc” indicates that there is no significant difference between “b” and “c”, but there is a significant difference between “a” and “ab”, and so on.
WA02	11.0c	7.01bc	7.67c	0.06bc	0.99b	30.73c	189.00ab	343.00aab
WA03	15.0ab	6.51bc	7.90b	0.06bc	0.75bc	37.97bc	131.00b	257.00bc
WA04	16.0a	26.77ab	7.90b	0.09b	1.02ab	39.93ab	134.33ab	274.00b
WA05	16.0a	12.61ab	7.80bc	0.26a	2.06a	42.87ab	132.00ab	289.33ab
WA06	16.0a	34.07a	7.90b	0.14ab	1.29ab	40.63ab	136.00ab	283.00ab
WA07	16.0a	13.01ab	7.90b	0.11ab	0.97bc	41.07ab	124.33bc	279.33ab
WA08	16.0a	8.05b	8.00ab	0.07bc	0.71bc	38.07bc	127.00bc	277.00ab
WA09	16.0a	12.71ab	8.07ab	0.13ab	0.69bc	39.63bc	144.00ab	273.00bc
WA10	15.0ab	6.25bc	8.10a	0.10ab	0.56c	39.70b	117.00bc	267.33bc
WA11	15.0ab	4.30c	8.10a	0.06c	0.77bc	35.80bc	115.33c	251.00c
WA12	14.5b	21.90ab	7.90b	0.10ab	1.13ab	70.60a	228.00a	465.00a

云南纳帕海高原湿地水体酵母菌多样性及其与理化因子的相关性

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况利沙 ¹^,² , 王海雁 ¹^,²^,³ , 雷洪涛 ¹^,² , 马跃强 ¹^,² , 韩荣琳 ¹^,² , 王留英 ¹^,² , 陈根 ¹^,² , 王永霞 ¹^,² , 赖泳红 ¹^,² , 李治滢 ¹^,²^,^*

微生物学报 | 土壤微生物与元素循环 2024,64(6): 1779-1799

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微生物学报 | 土壤微生物与元素循环 2024, 64(6): 1779-1799

云南纳帕海高原湿地水体酵母菌多样性及其与理化因子的相关性

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况利沙¹^,², 王海雁¹^,²^,³, 雷洪涛¹^,², 马跃强¹^,², 韩荣琳¹^,², 王留英¹^,², 陈根¹^,², 王永霞¹^,², 赖泳红¹^,², 李治滢¹^,²^,^*

作者信息

1 云南大学云南省微生物研究所, 云南昆明 650504

2 西南微生物多样性教育部重点实验室, 云南昆明 650504

3 玉林市林业科学研究所, 广西玉林 537501

Yeast diversity in water bodies and its correlations with physicochemical factors of Napahai plateau wetland, Yunnan Province

Lisha KUANG¹^,², Haiyan WANG¹^,²^,³, Hongtao LEI¹^,², Yueqiang MA¹^,², Ronglin HAN¹^,², Liuying WANG¹^,², Gen CHEN¹^,², Yongxia WANG¹^,², Yonghong LAI¹^,², Zhiying LI¹^,²^,^*

Affiliations

1 Yunnan Institute of Microbiology, Yunnan University, Kunming 650504, Yunnan, China

2 The Key Laboratory for Southwest Microbial Diversity of the Ministry of Education, Kunming 650504, Yunnan, China

3 Yulin Forestry Science Research Institute, Yulin 537501, Guangxi, China

出版时间: 2024-06-04 doi: 10.13343/j.cnki.wsxb.20230708

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摘要

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【目的】研究纳帕海水体酵母菌的种群多样性、空间分布特征及其与理化因子的相关性。【方法】采用醋酸纤维素滤膜过滤平置培养方法，分离纳帕海水体酵母菌；利用26S rRNA D1/D2区域序列分析和形态特征对分离得到的酵母菌进行分类鉴定；运用R 4.2.2和Canoco 5软件分析纳帕海水体酵母菌的多样性及其与理化因子的相关性。【结果】从纳帕海高原湿地水体中共分离获得565株酵母菌，鉴定为21个属，36个种。优势种为胶红酵母(Rhodotorula mucilaginosa)、季也蒙迈耶氏酵母(Meyerozyma guilliermondii)以及禾本红酵母(Rhodotorula graminis)。总氮是影响纳帕海高原湿地水体酵母菌种群数量的主要理化因子，与酵母菌种群数量呈显著性负相关。【结论】纳帕海水体酵母菌资源比较丰富且存在明显的空间异质性，人类活动和总氮对纳帕海水体酵母菌数量分布影响较大。纳帕海高原湿地水体酵母菌的群落组成与其他高原淡水湖泊具有较大差异，与其他高原水环境相比，纳帕海高原湿地具有独特的酵母菌资源，有待进一步研究。

关键词

纳帕海高原湿地 / 酵母菌 / 多样性 / 理化因子

Abstract

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[Objective] To study the population diversity, spatial distribution characteristics, and correlations with physicochemical factors of yeasts in the Napahai plateau wetland. [Methods] Yeast strains were isolated from the water samples of the Napahai plateau wetland by membrane filtration and plate culture. The strains were identified based on the D1/D2 sequences of the 26S rRNA gene and morphological characteristics. R 4.2.2 and Canoco 5 were used to analyze the yeast diversity and its correlations with physicochemical factors. [Results] A total of 565 yeast strains were isolated from the water bodies in the Napahai plateau wetland and identified as 36 species of 21 genera. The dominant species wereRhodotorula mucilaginosa,Meyerozyma guilliermondii, andRhodotorula graminis. Total nitrogen was the main physicochemical factor affecting the yeast count, which presented a negative correlation. [Conclusion] The water bodies of Napahai plateau wetland harbor abundant yeast resources which present significant spatial heterogeneity. Human activities and total nitrogen have significant impacts on the yeast count. The yeast community composition in the water bodies of the Napahai plateau wetland is significantly different from that in other plateau freshwater lakes and has unique yeast resources that require further research.

Key words

Napahai plateau wetland / yeasts / diversity / physicochemical factors

引用本文

况利沙, 王海雁, 雷洪涛, 马跃强, 韩荣琳, 王留英, 陈根, 王永霞, 赖泳红, 李治滢. 云南纳帕海高原湿地水体酵母菌多样性及其与理化因子的相关性. 微生物学报, 2024 , 64 (6) : 1779 -1799 . DOI: 10.13343/j.cnki.wsxb.20230708

Lisha KUANG, Haiyan WANG, Hongtao LEI, Yueqiang MA, Ronglin HAN, Liuying WANG, Gen CHEN, Yongxia WANG, Yonghong LAI, Zhiying LI. Yeast diversity in water bodies and its correlations with physicochemical factors of Napahai plateau wetland, Yunnan Province[J]. Acta Microbiologica Sinica, 2024 , 64 (6) : 1779 -1799 . DOI: 10.13343/j.cnki.wsxb.20230708

正文

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酵母菌广泛存在于自然界中，比如土壤、水体、植物^[1-2]、南极海洋^[3]、南极深海沉积物^[4]、南极土壤^[5]、北极冰川^[6]、南极冰川^[7]和岩石^[8]等。近年来，国内对于高原淡水酵母菌多样性的研究主要集中在云南^[9-17]和西藏^[18-19]，表明高原淡水湖泊酵母菌具有较高的研究价值。在国外，Russo等^[20]研究了阿根廷的Rio Agrio-Lake Caviahue火山酸性水体(pH 1.8−6.7)的酵母多样性；Brizzio等^[21]在阿根廷Patagonia西北部的冰川分离出担子菌酵母并对其细胞外酶活性进行了研究；Brandão等^[22]在巴西东南部的3个湖泊中研究了在37 ℃环境下生长的酵母菌的多样性、抗真菌性和抗药性；Branda等^[23]对意大利Calderone Glacier冰川中的酵母菌多样性进行了探究。

纳帕海湿地具有低纬度、高海拔、季节性、强紫外线和半封闭性等特点，是我国唯一独特的湿地，地处迪庆藏族自治州香格里拉市境内，坐标位于27°48′36″−27°54′27″N，99°37′12″−99°40′30″E，与青藏高原相连^[24]，于2005年被批准纳入到国际重要湿地名录。纳帕海高原湿地与周围具有丰富物种的森林植被形成湿地生态系统，由草甸、沼泽、水面和湖周森林构成的喀斯特型季节性沼泽湿地，孤立而分散，湿地之间没有水道相通。近年来，有研究者对纳帕海高原湿地微生物多样性及其与理化因子相关性进行研究^[25]，通过宏基因组研究纳帕海高原湿地微生物群落多样性，并对其固碳途径、氮代谢途径、病毒种群结构模式和病毒基因组中辅助代谢基因进行了初步分析^[26-27]，还有研究者对纳帕海高原湿地浮游病毒^[28]和噬菌体进行了相关研究^[29-30]。

本研究旨在探究纳帕海高原湿地水体酵母菌的结构组成、空间分布特征及其与理化因子的相关性；探究人类活动对纳帕海高原湿地酵母菌群落的影响以及酵母菌在该生态系统中扮演的角色，并为纳帕海高原湿地酵母菌作为纳帕海水体评价的潜在生物指示器提供参考。

1 材料与方法

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1.1 材料

1.1.1 样品采集

2022年6月15日在纳帕海27°48′36″−27°54′27″N，99°37′12″−99°40′30″E范围内沿着湖边采集11个水体样品，并记录每个样点的经纬度、海拔高度和生境信息(表1)。水样的采集选择采水器在水表面下深度约10−50 cm进行水样收集，每个水样点共采集5 L，分装成2份储存在2个无菌水桶中。1份用于酵母菌的分离，1份进行理化因子指标的测定。

1.1.2 主要仪器

Alpha^TM Unit Block Assembly for DNA Engine^® Systems，Bio-Rad公司；电热恒温鼓风干燥箱，上海跃进医疗器械有限公司(HENGZI)；灭菌锅，SANYO公司；隔水式电热恒温培养箱，上海新诺仪器集团有限公司；小型高速离心机，ThermoFisher Scientific公司；小型电泳仪，Bio-Rad公司；凝胶成像仪，Syngene公司。

1.1.3 培养基

培养基的配制参考李治滢等^[13]和周新丽等^[14]的方法，并稍作改动(将培养基中的水换成纳帕海水)。

1.2 酵母菌的分离

使用孔径0.45 μm，直径50 mm的醋酸纤维素滤膜对水样进行过滤，每个水样过滤36次，每次过滤50 mL，共过滤1 800 mL。分别平置于12个酸化YM培养基、含氯霉素的RBCH培养基、酸化马铃薯葡萄糖琼脂(potato dextrose agar, PDA)培养基平皿上，并置于室温培养。培养2−4 d后计算酵母菌的数量，挑取生长的酵母菌于常规YM平板培养基上进行划线纯化培养，得到的酵母菌转接到YM斜面保存留用。酵母菌数量CFU (个/L)=(酵母菌数量平均值/过滤水量)×1 L。

1.3 形态观察、DNA的提取和26S rRNA D1/D2区域的扩增

对分离获得的酵母菌进行形态观察^[31]、DNA的提取和26S rRNA D1/D2区域的扩增^[32-33]。扩增引物序列：NL1 (5′-GCATATCAATAAGCGGAGGAAAAG-3′)和NL4 (5′-GGTCCGTGTTTCAAGACGG-3′)。扩增产物送至生工生物工程(上海)股份有限公司进行测序。

1.4 鉴定

结合酵母菌的形态特征和26S rRNA D1/D2区域序列分析和构建的系统发育树，参照文献[31,34-35]对分离获得的酵母菌进行分类鉴定。

1.5 序列分析和系统发育树的构建

对酵母菌的26S rRNA D1/D2区域进行序列分析^[36]。采用DNAStar软件对酵母测序所得到的26S rRNA D1/D2序列进行人工校对，在GenBank核酸序列数据库中进行同源序列搜索(BLAST search)，采用MEGA 11.0软件的邻接法进行系统发育树分析^[37-39]，使用Kimuran 2-parameter distance模型及neighbour-joining方法构建系统发育树，boostrap值的计算分析根据1 000次随机抽样进行，得到酵母菌系统进化树。

1.6 理化指标测定

测定水样的pH、温度(temperature, Temp)、浊度(turbidity, Turb)、总磷(total phosphorus, TP)、总氮(total nitrogen, TN)、总有机碳(total organic carbon, TOC)、总硬度(total hardness, TH)和电导率(electrical conductivity, EC)等指标。温度指标使用温度计在采样现场进行测定；浊度指标使用浊度仪(Eutech Instruments Pte Ltd.)在实验室进行测定，重复3次并取其平均值。其余指标委托“昆明绿岛环境科技有限公司”检测。

1.7 数据分析

采用出现频率(frequency of occurrence, OF)比较酵母菌在各样点分布，OF=酵母菌种群出现的样点数/总样点数×100；分离频率(IF)比较判断酵母菌的优势类群，IF=某一酵母菌种群的菌株获得数/分离到的总酵母菌的菌株数×100。通过生物统计原理计算酵母菌的Simpson index、Shannon-Weiner index、Richness evenness、Pielou evenness、Chao1 value。利用R 4.2.2进行jaccard相似度指数和Spearman相关性系数的分析以及绘制相关图片。使用Canoco 5软件对酵母菌类群分布与环境理化因子进行主成分分析(principal component analysis, PCA)。

2 结果与分析

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2.1 纳帕海高原湿地水体酵母菌的多样性

2.1.1 纳帕海高原湿地水体酵母菌的种群组成

从纳帕海高原湿地的11个水体样品中共分离获得565株酵母菌。将已鉴定的酵母菌26S rRNA D1/D2区域序列上传GenBank获取菌株登录号(OR083772−OR083835，OR917881−OR917889)。结合酵母菌的形态特征和26S rRNA D1/D2区域的序列分析，将分离获得的酵母菌分类鉴定为21个属，36个种(表2)，并对CFU进行了统计。选择36个种的代表菌株与相应模式菌株的26S rRNA D1/D2区域序列构建系统进化树。

如图1所示，纳帕海高原湿地水体酵母菌由子囊菌酵母和担子菌酵母两大类群组成。由表2可知，水体中的子囊菌酵母共有166株，归8个属12个种；担子菌酵母有399株，归13个属24个种；未分离获得类酵母。在纳帕海高原湿地所有水体样点中均分离到子囊菌酵母和担子菌酵母，其中担子菌酵母为主要类群。

在纳帕海高原湿地水体酵母菌的种群结构中，Naganishia属有4个种；假丝酵母属(Candida)、维希尼克氏酵母属(Vishniacozyma)和红酵母属(Rhodotorula)分别有3个种；耶氏酵母属(Yarrowia)、地霉属(Geotrichum)、黑粉菌属(Ustilago)、Papiliotrema、囊担菌属(Cystobasidium)以及线黑粉酵母属(Filobasidium)分别得到2个种；迈耶氏酵母属(Meyerozyma)、Kurtzmaniella以及路德酵母属(Lodderomyces)等共11个属仅分别分离得到1个种。其中Rhodotorula属的3个种分离株数高达261株，分离频率(IF)为46.20%，为纳帕海高原湿地水体酵母菌优势属；其次是Meyerozyma属，该属仅有1种，但分离株数达到102株，IF为18.05%。纳帕海高原湿地水体酵母菌的优势种为胶红酵母(Rhodotorula mucilaginosa) (IF=33.63%)、季也蒙迈耶氏酵母(Meyerozyma guilliermondii) (IF=18.05%)和禾本红酵母(Rhodotorula graminis) (IF=12.04%)为；解脂耶罗维亚酵母(Yarrowia lipolytica)、维多利亚维希尼克氏酵母(Vishniacozyma victoriae)、威林格氏线黑粉菌(Filobasidium wieringae)等9个种分别仅得到1株，IF仅分别为0.18%。分析其出现频率可知，Rhodotorula mucilaginosa出现频率最高(OF=100.00%)，所有样点均有出现；其次为Rhodotorula graminis、Meyerozyma guilliermondii以及Meira nashicola的OF分别为90.91%、72.73%、54.55%；而微黄汉纳酵母(Hannaella luteola)、Vishniacozyma victoriae、Cystobasidium lysinophilum等14个种仅在1个样点被分离得到，OF分别为9.10%。在纳帕海高原湿地水体酵母菌数量中，纳曲河入河口WA08样点酵母菌数量最多，CFU为8 472个/L；其次是靠近人为居住地的死水处WA09样点，CFU为8 372个/L；而人员居住少、远离人员居住地和水中植物少的WA05、WA06和WA04样点的CFU少，分别为200、615和700个/L。

2.1.2 纳帕海高原湿地酵母菌的相似性分析

对纳帕海高原湿地各个水体样点酵母菌的Jaccard相似性进行了分析(表3)。从表3数据得知，纳帕海高原湿地水体酵母菌的种群相似度系数范围0.05−0.57；同为旅游区的WA02样点和WA03样点达到中等相似(0.50−0.75)，占比为1.82%；有大量游客的WA02样点和远离人员居住地的WA04、WA10样点等22组样点间相似度为中度不相似(0.25−0.50)，比例为40.00%；32组样点为极度不相似(0.00−0.25)，比例为58.18%，其中有3组样点酵母菌Jaccard相似度系数范围为0.00−0.10。总而言之，纳帕海高原湿地水体各样点酵母菌在种水平上分布存在较大的差异。

WA02样点和WA03样点达到中度相似，这两个样点都是处于旅游区附近，受到较多的人为活动干扰。所有样点间均未出现相似度为0的情况，远离人员居住地且临山的WA04和靠近人员居住地且水中植物较多的WA12的相似度最低，为0.05，这两个样点间不仅距离较远，且生境也相差颇大。除此之外，WA12样点与每一个样点的相似度皆为最低，WA12号样点距离所有样点都比较远，且WA12样点具有独特的水体生境，较多的水生植物且大部分为浮水植物。然而，相较于其他样点，以挺水植物为主的WA11样点同样具有较多的水生植物，因此差异性较小。

2.1.3 纳帕海高原湿地酵母菌多样性指数分析

使用R 4.2.2计算纳帕海高原湿地水体酵母菌的多样性指数。如图2所示，WA04、WA03和WA09样点总丰度最高，其次是WA07和WA11样点。多样性指数表明，WA04样点Shannon-Wiener多样性指数最高，其次是WA09、WA03、WA11、WA07和WA02样点，WA10和WA12样点Shannon-Wiener多样性指数最低；Simpson多样性指数分析结果中，WA04样点指数最低，其次是WA07、WA09、WA11、WA03以及WA02样点的偏低，WA10和WA12样点的最高。可以进一步观察得出，Shannon-Wiener多样性指数和Simpson多样性指数趋势相反，Shannon-Wiener和Richness丰富度指数以及Chao1多样性指数趋势基本一致。这4个指数综合分析表明，在纳帕海高原湿地水体中，远离人类活动的山区WA04样点分离得到的酵母菌丰富度和多样性最高，人为活动干扰较大的WA03、人为干扰较小的天然松林WA07、人为干扰较大的死水区WA09以及水生植物丰富且挺水植物居多的WA11样点丰富度和多样性偏高，位于乔木灌木林旁边的WA10和水生植物丰富且浮水植物居多的WA12样点分离得到的酵母菌丰富度和多样性最低。Pielou均匀度指数的趋势则不相同，在纳帕海高原湿地水体中，WA12号样点分离到的酵母菌菌株分布最均匀，其次是WA07、WA11和WA04样点，而WA10样点中分离到的酵母菌菌株则最不均匀。

2.2 纳帕海高原湿地酵母菌多样性与理化因子的相关性

2.2.1 纳帕海高原湿地水体理化因子差异性分析

为揭示纳帕海高原湿地酵母菌的多样性和空间分布特征及其与理化因子之间的相关性，探讨人类活动对纳帕海高原湿地酵母菌的影响以及酵母菌在该生态系统中扮演的角色。测定了纳帕海高原湿地水体8个理化指标数据(表4)。

由表4可以看出，纳帕海高原湿地11个水样中，理化因子存在不同程度的差异。各个样点的水温为11.0−16.0 ℃，其中WA02样点水温最低，为11.0 ℃，WA04−WA09样点温度最高，均为16.0 ℃，除WA02样点与WA12样点，其余样点间均不存在显著差异性(P > 0.05)；水体浊度为4.30−34.07，具有较多挺水植物且远离人员居住地的WA11样点最低，为4.30，位于人工林旁WA06样点最高，为34.07；pH为7.67−8.10，其中WA02样点最低，为7.67，WA10和WA11样点最高，均为8.10；TP为0.06−0.26 mg/L，其中具有较多挺水植物且远离人员居住地的WA11样点最低，为0.06 mg/L，水中有泡沫的WA05样点最高，为0.26 mg/L；TN为0.56−2.06 mg/L，其中有山泉水冒出的WA10样点最低，为0.56 mg/L，水中有泡沫的WA05样点最高，为2.06 mg/L；TOC为30.73−70.60 mg/L，其中无植物生长WA02样点最低，为30.73 mg/L，靠近人员聚集地具有较多的水生植物WA12样点最高，为70.60 mg/L；TH为115.33−228.00 mg/L，其中具有较多挺水植物且远离人员居住地的WA11样点最低，为115.33 mg/L，靠近人员聚集地具有较多的水生植物WA12样点最高，为228.00 mg/L；EC为251.00−465.00 μS/cm，具有较多挺水植物且远离人员居住地的WA11样点最低，为251.00 μS/cm，靠近人员聚集地具有较多的水生植物的WA12样点最高，为465.00 μS/cm。

2.2.2 纳帕海高原湿地水体样品酵母菌多样性与理化因子之间的相关性分析

为探究纳帕海高原湿地水体酵母菌的多样性和空间分布特征及其与理化因子之间的相关性，对纳帕海高原湿地11个水样的CFU、物种数(species)、Shannon-Winner多样性指数、Simpson多样性指数、Pielou均匀度指数、Richness丰富度指数、Chao1指数以及理化因子进行Spearman相关性分析(图3)。物种数与Shannon-Winner多样性指数、Richness丰富度指数、Chao1指数均存在极其显著的正相关性(极其显著P < 0.001，下同)，与Simpson多样性指数呈极其显著负相关，其余多样性指数之间也存在一定的关联性，这与多样性分析的结果一致。CFU与总氮呈现显著负相关(显著P < 0.05，下同)，相关系数为−0.59；浊度与总磷以及总有机碳之间呈现极显著正相关(极显著P < 0.01，下同)，相关系数分别为0.63、0.67，与总氮之间呈现显著正相关，相关系数为0.62；pH与总氮以及电导率之间呈现极其显著负相关，相关指数分别为−0.73和−0.70；总磷和总有机碳之间呈现极其显著正相关，相关系数为0.79，与浊度和温度都呈现出正相关关系；电导率与总氮以及总硬度之间呈现极显著正相关，相关系数分别为0.65和0.66，与pH之间呈现负相关；其他参数间则没有明显相关性。酵母菌多样性和理化因子之间的Spearman相关系数表明，Species、Shannon-Winner多样性指数以及Richness丰富度指数与总磷、总有机碳、总氮和电导率之间都呈现出负相关关系。有研究表明，水质越清洁多样性越丰富^[40]，有机质含量越多，富营养化程度越重，越污染，多样性越少^[41]。这与本次实验结果一致。Spearman相关性分析表明，酵母菌数量与总氮之间呈现负相关，总氮会影响纳帕海水体酵母菌的数量，多样性与环境理化因子之间呈现负相关。

2.2.3 纳帕海高原湿地酵母菌多样性与理化因子之间的PCA分析

使用Canoco 5软件进行PCA分析。确定合适的响应模型之前，对采样点和数据进行去趋势对应分析(detrended correspondence analysis, DCA)。如图4A所示，对纳帕海高原湿地水体样点多样性与理化因子之间进行DCA分析，显示第一排序轴的梯度长度值大小为0.39，该值小于4.00，选用非线性排序中的PCA来分析，结果显示第一主成分对响应变量的解释度是58.22%，第二主成分对响应变量的解释度为86.58%。如图4B所示，对纳帕海高原湿地水体样点种群与理化因子之间进行DCA分析，显示第一排序轴的梯度长度值大小为1.27，该值小于4.00，选用非线性排序中的PCA来分析，结果显示第一主成分对响应变量的解释度是49.88%，第二主成分对响应变量的解释度为69.64%。

如图4A所示，其样点分布状况显示，WA02、WA03、WA07、WA08、WA09以及WA11样点排序比较接近；WA05和WA06样点聚在一起；WA10以及WA12样点聚在一起；而WA04独占一方，距离较远。结合表2，观察纳帕海高原湿地酵母菌种群在各样点的分布情况。Rhodotorula mucilaginosa和Rhodotorula graminis几乎在每个样点都有出现。Yarrowia lipolytica、甜茅黑粉菌(Ustilago filiformis)、Ustilago longissima、Cystofilobasidium infirmominiatum、Vishniacozyma taibaiensis、卡恩斯维希尼克氏酵母(Vishniacozyma carnescens)、Vishniacozyma victoriae、阿德利长西氏酵母(Naganishia adeliensis)和Cystobasidium lysinophilum仅在WA04样点出现，WA04样点出现足有9个特有种，导致WA04样点独自占有一个区域。WA10与WA12样点所拥有的酵母菌种群最少，WA12样点仅拥有2个不同的酵母菌种群，而两个样点共同拥有长孢洛德酵母(Lodderomyces elongisporus)和Rhodotorula mucilaginosa，因此WA10以及WA12样点聚在一起。Filobasidium wieringae只出现在WA05样点，Geotrichum bryndzae在WA06样点中大量分离，且WA05和WA06样点共同分离得到Meyerozyma guilliermondii、Meira nashicola、胶红酵母(Rhodotorula mucilaginosa)以及禾本红酵母(Rhodotorula graminis)，因此WA05和WA06样点聚在一起且相互有一定的距离。Meyerozyma guilliermondii和Papiliotrema aspenensis大量在WA02、WA03、WA07、WA08、WA09以及WA11样点中分离得到；Kurtzmaniella quercitrusa、热带假丝酵母(Candida tropicalis)、Barnettozyma californica、Yarrowia osloensis、蚜虫莫氏黑粉菌(Moesziomyces aphidis)、Papiliotrema flavescens、Naganishia diffluens、Apiotrichum laibachii和圆红酵母(Rhodotorula toruloides)仅在WA02、WA03、WA07、WA08、WA09以及WA11样点中出现；而涎沫假丝酵母菌(Candida zeylanoides)仅在WA02样点出现，Hannaella luteola仅在WA07样点出现，Naganishia cerealis仅在WA11样点出现，这些条件导致WA02、WA03、WA07、WA08、WA09以及WA11样点独立于其他样点却相互之间互相分离。分析其相关性，如果它们之间的夹角等于90°则没有相关性，小于90°则呈正相关关系，反之则呈现负相关关系。酵母菌数量与总氮呈现显著负相关，与图4的结果一致。物种数与Shannon-Winner多样性指数、Simpson多样性指数、Richness丰富度指数、Chao1指数均存在极其显著的正相关性，这也与之前的结果一致。

为了进一步观察理化因子对酵母菌种群数量的影响，将数量较多的酵母菌种群数量与理化因子之间进行PCA分析，如图4B所示。总硬度、电导率、总有机碳与N.albida、L.elongisporus、R.mucilaginosa呈现出正相关关系，与M.aphidis、P.aspenensis、M.guilliermondii、R.graminis和P.flavescens等呈现负相关关系，总硬度、电导率、总有机碳与样点的酵母菌数量并无明显关系。浊度、总磷只与M.nashicola和R.graminis呈现正相关，与L.elongisporus、R.mucilaginosa、P.flavescens等呈现负相关，与样点的酵母菌数量呈现负相关。总氮只与M.nashicola呈现出正相关关系，与其余大部分酵母菌种群都呈现负相关，包括数量排名最多的几个酵母菌种群M.guilliermondii、R.graminis和R.mucilaginosa，因此总氮能显著影响纳帕海高原湿地酵母菌种群数量。总而言之，总氮是影响纳帕海高原湿地酵母菌种群分布最重要的理化因子，主要通过影响大部分酵母菌种群的数量。

2.2.4 纳帕海高原湿地酵母菌多样性与其他淡水环境之间的分析

分析比较纳帕海湿地与云南高原湖泊滇池^[9]、抚仙湖^[12]、星云湖^[40]、杞麓湖^[11]、程海^[13-14]、阳宗海^[10,15]、洱海^[15]、异龙湖^[16-17]和西藏自治区的拉鲁湿地^[19]、羊卓雍措^[18]的酵母菌种群关系。结果发现，胶红酵母(Rhodotorula mucilaginosa)是所有样点均有的一个酵母菌种群。再对其相似性进行分析(图5)，纳帕海高原湿地酵母菌种群与其他高原水环境中酵母菌种群具有明显差异，物种相似度为0.055 6–0.472 2，共有种为1–17个。相比之下，纳帕海高原湿地与滇池(DL)酵母菌种群的相似度最高为0.472 2，水质均为Ⅴ类左右，同时又具有水质稍微好一点的区域；纳帕海是属于Ⅴ类水质，与异龙湖湿地修复前相似度第二高为0.277 8，而与异龙湖湿地修复后相似度有所降低；整体来看，纳帕海高原湿地与其他几个湿地之间的相似度都是略高于其他淡水湖泊。纳帕海高原湿地与其他高原淡水湖泊之间的相似性均不相同，这可能与其地理环境和水质不同有关。

纳帕海高原湿地独有的酵母菌种群有Barnettozyma californica、Candida pseudointermedia、Cutaneotrichosporon dermatis、Cystobasidium lysinophilum、Kurtzmaniella quercitrusa、Meira nashicola、Naganishia cerealis、Ustilago filiformis、Ustilago longissima、Vishniacozyma victoriae、Yarrowia osloensis。这可能与纳帕海高原湿地具有独特的地理位置、气候以及生态环境有关。

3 讨论

收起

3.1 纳帕海高原湿地多样性

研究表明纳帕海高原湿地水体酵母菌具有丰富的多样性。本研究共采集水体样品11个，分离获得565株酵母菌，鉴定为21个属，36个种。优势种为胶红酵母(Rhodotorula mucilaginosa)、季也蒙迈耶氏酵母(Meyerozyma guilliermondii)以及禾本红酵母(Rhodotorula graminis)；优势属为红酵母属(Rhodotorula)。红酵母属广泛存在于自然界中且抗逆性较强，特别是在高海拔水环境中，无论水质与生态环境是否相同，红酵母属都广泛存在。另外，在中国云南星云湖^[42]、中国青藏高原的雪坑^[43]、中国新疆的天山一号冰川^[44-45]、阿根廷的Nahuel Huapi湖^[46]、阿根廷巴塔哥尼亚的Rio Agrio和Caviahue湖^[20]、阿根廷巴塔哥尼亚的冰川^[47]、意大利的Calderone Glacier冰川^[23]以及巴西Lagoa Santa高原的Lagoa Olhos d’Agua湖^[22]都分布有红酵母属的酵母菌。这些环境都具有高海拔和强紫外线的共同特征，因为红酵母属能够产生类胡萝卜素等色素，而这些色素具有优秀的紫外线辐射抵抗能力，使酵母菌能够更好地适应强紫外线辐射环境，在高海拔环境中占据优势。

Meyerozyma guilliermondii是纳帕海高原湿地中的另外一种优势菌。M.guilliermondii是一种子囊菌酵母，可以从环境、人类皮肤以及黏膜中(作为一种腐生菌)分离得到。Gadanho等^[48]从大西洋的深海热液区分离得到，ChoI等^[49]从泡菜生产过程中产生的废盐水环境中分离到1株。M.guilliermondii不产色素，在橄榄油培养基中能产生脂肪酶，在海水培养基中可以产生大量菊粉酶，它被认为是海洋中的土著种，在海洋环境中广泛分布^[50]，除此之外，M.guilliermondii还能合成核黄素^[51]。有研究发现M.guilliermondii是一种罕见的病原菌，可引起心包炎、心内膜炎、腹膜炎、骨髓炎和导管介导的念珠菌病等^[52]。又有研究发现M.guilliermondii具有抗菌特性，通过在水果的伤口表面定居并争夺营养物质和生态位的方式拮抗水果病原菌^[53]。从梨表面分离的M.guilliermondii已成功应用于几种水果和蔬菜采后病原体的防控，如葡萄、苹果以及番茄等^[54]。然而在淡水环境中，M.guilliermondii并不常见。星云湖(Ⅴ类)分离得到4株，异龙湖(Ⅴ类)中分离得到2株，杞麓湖(Ⅴ类)中分离得到2株，阳宗海(Ⅲ类)分离得到4株，这4个淡水湖泊普遍水质较差。由此可见，在淡水环境中，M.guilliermondii普遍生存在水质恶劣的环境中，M.guilliermondii具有对抗恶劣环境的可能，同时该菌作为一种潜在的病原菌，也在一定程度上说明了环境中可能存在有致病风险。在以前报道的其他淡水环境中，M.guilliermondii分离频率都较低，而本次的研究结果M.guilliermondii首次作为优势种群出现在淡水环境中，应该是纳帕海高原湿地独特的气候与生态环境所造成，也说明了纳帕海环境保护是目前需要重点关注的问题。

在纳帕海高原湿地水体中，WA04样点受到人为干扰较小，分离得到的酵母菌多样性最高；人为活动干扰较大的WA02、WA03和死水区WA09样点多样性偏高；人为活动干扰较小的WA10样点多样性偏低，有5种；人为活动干扰较大且浮水植物居多的WA12样点分离得到的酵母菌多样性最低，仅有2种。人为活动等因素并不能很好地与酵母菌多样性相关联。挺水植物对氮磷具有较好的去除能力，由此挺水植物居多的WA11具有最低的总磷和较低的总氮，但与酵母菌的多样性未观察到相关性。而浮水植物对于氮磷的去除能力相比挺水植物更弱一些，WA12样点总氮和总磷比WA11要稍微高一些，但WA12样点总有机碳、总硬度以及电导率都是所有样点中最高，较高的有机物导致浮水植物大量繁殖，水体中溶氧不足，水质恶化，因此WA12样点酵母菌多样性最低。

3.2 纳帕海高原湿地酵母菌数量分布与人类活动相关性

纳帕海高原湿地作为云南省著名旅游区，围绕纳帕海建造了环纳帕海公路，公路与水体接壤处堆积大量石子防止泥土被冲刷到公路上，而本次采样有好几个水体样点位于公路边。WA05样点水体紧靠公路边且靠近人员居住地，水中有明显泡沫，水体酵母菌数量最少。样点WA04和WA06同样位于公路边，但远离人类居住地，无明显水生植物生长，因此该样点酵母菌数量较少，但比样点WA05稍多。WA11样点离公路距离稍远且远离人员居住地，位于灌木林旁边，水生植物丰富且挺水植物居多，相比于WA04、WA05和WA06三个样点，WA11样点酵母菌数量突破1 000个/L，达到2 529个/L。样点WA10相比于WA11样点距离公路同样稍远且远离人类居住地，但位于乔木林旁边，水中具有挺水植物，水边有大量天然的草本植物，酵母菌数量大于3 000个/L，比WA11样点酵母菌数量稍多。按照水质情况来讨论，WA02、WA03和WA08样点总磷达到Ⅱ类水质标准，总氮达到了Ⅲ类水质标准。WA02和WA03样点位于旅游区，且WA02样点有大量马匹停留，WA02样点酵母菌数量有3 100个/L，WA03样点有3 800个/L，而WA08样点位于纳曲河森林入河口，远离人类居住地，酵母菌数量为8 472个/L，是WA02和WA03样点的2倍多。WA07和WA09样点总磷、总氮都达到了Ⅲ类水质，均远离人类居住地，但WA07样点紧靠公路且旁边为天然松林，WA09样点位于死水区且水岸有大量天然的野草以及少量乔本植物，WA07样点的酵母菌数量比WA09样点稍微少一些。说明相似的水质条件下，人为干扰会极大地影响酵母菌的数量分布。同样是受人为因素干扰较多和总氮为Ⅳ类水质以下的样点WA04、WA05、WA06和WA12，WA12样点由于其水生植物丰富，酵母菌数量显著比另外3个样点多。说明相似的水质以及受人为因素的干扰，水生生态环境也会影响其酵母菌的数量分布。由此可以得出，水质越差，水体酵母菌数量越少；随着远离公路的距离增加，酵母菌数量也会逐渐增加；水体旁边的植被也会直接影响到水体酵母菌的数量，具体表现为植被干扰程度越小，水体酵母菌数量越多；而水体中的植被生长情况也会影响到酵母菌数量，有水生植物生长的水体，酵母菌数量会多于无水生植物生长的水体。

3.3 理化因子相关性

3.3.1 纳帕海高原湿地理化因子

理化因子差异性分析结果表明纳帕海水体中不同样点之间的理化因子差异性明显。造成差异的原因是湿地环境内水体流动较少，不能及时进行水体交换，再加上不同区域的植被类型、土壤类型、水生生态以及人员活动等因素，极大程度上影响了纳帕海高原湿地水体的理化因子，导致酵母菌群落组成的差异。根据中华人民共和国生态环境部发布的地表水环境质量标准，纳帕海水体总氮在Ⅲ−劣Ⅴ类，总磷在Ⅱ−Ⅲ类，明显可以看出纳帕海高原湿地污染类型主要是氮污染。由于旅游业以及畜牧业的开发，纳帕海生态环境受到大量人为因素干扰，水体生态环境愈加恶劣，造成水体富营养化逐渐加重，纳帕海生态环境保护迫在眉睫。

3.3.2 影响纳帕海高原湿地与其他淡水环境酵母菌种群分布的环境因子

总氮是影响纳帕海酵母菌数量的主要因素，总氮与纳帕海酵母菌数量呈现显著性负相关，这与其他高原淡水酵母菌种群与理化因子之间的相关性分析并不相同。总磷和pH分别是影响杞麓湖中红冬孢酵母属(Rhodosporidium)和隐球酵母属(Cryptococcus)分布的主要因素；影响异龙湖酵母菌种群分布的是总氮；影响抚仙湖酵母菌种群的则是总有机碳；电导率是影响阳宗海酵母菌种群分布的主要因素；影响羊卓雍措酵母菌种群分布的主要因素是总磷和pH；化学需氧量则影响西藏拉鲁湿地水体酵母菌种群的分布；总氮和总磷影响巴西Rio Doce河流酵母菌属水平的分布^[55]。

这可能是由于各个环境内的地理位置、生态情况、气候差异以及人为活动都有所不同，导致酵母菌种群分布在不同环境中都有所差异，甚至于影响酵母菌种群分布的理化因子都有所不同。总氮作为衡量水质的重要指标之一，纳帕海高原湿地周边排放的生活污水和农田排水是主要影响纳帕海水体总氮的因素。再加上纳帕海高原湿地的绕湖公路紧紧围绕纳帕海水体建造，致使纳帕海水体受到人为因素干扰极大，几乎所有的样点都受到一定的干扰。因此，与人类活动息息相关的总氮便成为纳帕海水体影响酵母菌分布的关键因素。而其他淡水环境与纳帕海高原湿地环境不同，因此影响酵母菌分布的关键因素也有所不同。

3.4 纳帕海高原湿地多样性与其他高原淡水的比较

在纳帕海高原湿地与其他高原淡水环境酵母菌种群比较发现，Barnettozyma californica、Candida pseudointermedia、皮肤皮状新丝孢酵母(Cutaneotrichosporon dermatis)、Cystobasidium lysinophilum、Kurtzmaniella quercitrusa、Meira nashicola、Naganishia cerealis、Ustilago filiformis、Ustilago longissima、Vishniacozyma victoriae和Yarrowia osloensis都是纳帕海高原湿地独有的酵母菌种群。Naganishia cerealis、Ustilago filiformis、Ustilago longissima和Vishniacozyma victoriae这4种酵母菌分别只分离得到1株，可能是雨水或空气流动等原因带到水体中的，具有偶然性。Barnettozyma californica在铵态氮、硝酸盐或亚硝酸盐浓度为14−140 mg/L时表现出高效的异养硝化和好氧反硝化功能，具有从废水中去除无机氮的潜力^[56]。Candida pseudointermedia能将纤维二糖转运到细胞质中，然后在细胞内产生高亲和力的β-葡萄糖苷酶水解纤维二糖^[57]，同时，C.pseudointermedia也具有利用降解脂肪族和芳香烃作为营养物质来源的能力，其在油污染环境中的生物修复有效性，可以为水体或土壤油污染提供解决方案^[58]。Cutaneotrichosporon dermatis是一种能将木质纤维素转化为微生物脂质的含油酵母^[59]，能够同时吸收木糖和葡萄糖，从而产生相当大的脂质积累^[60]，C.dermatis对噻康唑、特比萘芬、氟康唑和泊沙康唑等都具有一定的耐药性^[61]，可有效降解四环素、多西环素和四氯环素等，从而降低抗真菌药的抗菌效力^[62]，是一种潜在致病菌，能产生一些重要的毒力因子^[63]。这些纳帕海水体中独有的酵母菌种群都会间接或直接影响到水质环境，构成了纳帕海高原湿地独特的生态环境。

4 结论

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本研究探讨了纳帕海高原湿地水体酵母菌多样性及其与理化因子的相关性。结果显示，纳帕海高原湿地水体中酵母菌资源丰富，共分离得到565株酵母菌，鉴定为21个属，36个种。人为活动对纳帕海高原湿地水体酵母菌的数量和种群分布影响较大。总氮是影响纳帕海高原湿地酵母菌种群数量的主要理化因子，总氮与酵母菌种群数量呈现显著性负相关。纳帕海高原湿地酵母菌群落组成与其他高原淡水湖泊具有较大差异，与其他高原淡水环境相比，纳帕海高原湿地具有独特的酵母菌资源，可作为工业领域的待开发菌种资源，有待进一步研究。

基金

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国家自然科学基金(31160006)
云南省重大科技专项(生物资源数字化开发应用)(202002AA100007)

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2024年第64卷第6期

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doi: 10.13343/j.cnki.wsxb.20230708

接收时间：2023-11-19
首发时间：2026-03-19
出版时间：2024-06-04

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收稿日期：2023-11-19
录用日期：2024-02-04

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National Natural Science Foundation of China(31160006)

国家自然科学基金(31160006)

Major Science and Technology Special Project of Yunnan Province (Digital Development and Application of Biological Resources)(202002AA100007)

云南省重大科技专项(生物资源数字化开发应用)(202002AA100007)

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1 云南大学云南省微生物研究所, 云南昆明 650504

2 西南微生物多样性教育部重点实验室, 云南昆明 650504

3 玉林市林业科学研究所, 广西玉林 537501

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2种不同金属材料的力学参数

科 Family	属数 Number of genus	种数 Number of species	占总种数比例 Percentage of total species (%)	属 Genus	种数 Number of species	占总种数比例 Percentage of total species (%)
鹅膏菌科Amanitaceae	2	11	5.26	鹅膏菌属 Amanita	10	4.78
小菇科 Mycenaceae	2	12	5.74	丝盖伞属 Inocybe	5	2.39
多孔菌科 Polyporaceae	8	14	6.70	蜡蘑属 Laccaria	5	2.39
红菇科 Russulaceae	3	23	11.00	小皮伞属 Marasmius	6	2.87
				小菇属 Mycena	11	5.26
				光柄菇属 Pluteus	5	2.39
				红菇属 Russula	17	8.13
				栓菌属 Trametes	5	2.39

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Sites	Longitude	Latitude	Altitude (m)	Overview of the surrounding environment of the sampling point
WA02	99°39′49″E	27°52′55″N	3 283.30	It’s a tourist area and a racecourse, with a large number of tourists
WA03	99°38′27″E	27°53′48″N	3 273.20	It’s a tourist area
WA04	99°38′9″E	27°54′8″N	3 284.60	It’s a mountainous area, and far from where people live
WA05	99°37′52″E	27°54′26″N	3 285.80	It’s a place where few people live, and there is foam in the water
WA06	99°37′43″E	27°54′18″N	3 285.69	It’s a planted forest, and far from where people live
WA07	99°37′48″E	27°53′50″N	3 274.55	It’s a natural pine forest, and far from where people live
WA08	99°37′43″E	27°53′36″N	3 272.30	It’s the mouth of the Naqu River, and far from where people live
WA09	99°37′26″E	27°53′53″N	3 272.30	It’s the cape of Napahai, with poor water mobility and mostly stagnant water, and there are people living nearby
WA10	99°37′17″E	27°53′8″N	3 272.30	It’s a spring hole in the water, with mountain spring water emerging, and far from the place where people live
WA11	99°37′52″E	27°51′43″N	3 272.30	There are many emergent plants in the water, and far from where people live
WA12	99°38′4″E	27°49′59″N	3 283.30	It’s a place with a large number of aquatic plants, and close to where people gather

Sites

Longitude

Latitude

Altitude (m)

Overview of the surrounding environment of the sampling point

WA02

99°39′49″E

27°52′55″N

3 283.30

It’s a tourist area and a racecourse, with a large number of tourists

WA03

99°38′27″E

27°53′48″N

3 273.20

It’s a tourist area

WA04

99°38′9″E

27°54′8″N

3 284.60

It’s a mountainous area, and far from where people live

WA05

99°37′52″E

27°54′26″N

3 285.80

It’s a place where few people live, and there is foam in the water

WA06

99°37′43″E

27°54′18″N

3 285.69

It’s a planted forest, and far from where people live

WA07

99°37′48″E

27°53′50″N

3 274.55

It’s a natural pine forest, and far from where people live

WA08

99°37′43″E

27°53′36″N

3 272.30

It’s the mouth of the Naqu River, and far from where people live

WA09

99°37′26″E

27°53′53″N

3 272.30

It’s the cape of Napahai, with poor water mobility and mostly stagnant water, and there are people living nearby

WA10

99°37′17″E

27°53′8″N

3 272.30

It’s a spring hole in the water, with mountain spring water emerging, and far from the place where people live

WA11

99°37′52″E

27°51′43″N

3 272.30

There are many emergent plants in the water, and far from where people live

WA12

99°38′4″E

27°49′59″N

3 283.30

It’s a place with a large number of aquatic plants, and close to where people gather

Species

Sampling sites

Subtotal

OF (%)

IF (%)

WA02

WA03

WA04

WA05

WA06

WA07

WA08

WA09

WA10

WA11

WA12

Ascomycetous

Barnettozyma californica

18.18

0.35

Candida pseudointermedia

27.27

0.53

Candida tropicalis

18.18

0.53

Candida zeylanoides

9.10

1.06

Debaryomyces fabryi

9.10

0.18

Geotrichum bryndzae

18.18

1.24

Geotrichum silvicola

27.27

0.71

Kurtzmaniella quercitrusa

18.18

0.53

Lodderomyces elongisporus

36.36

3.72

Meyerozyma guilliermondii

102

72.73

18.05

Yarrowia lipolytica

9.10

0.18

Yarrowia osloensis

18.18

1.95

Basidiomycetous

Apiotrichum laibachii

18.18

1.06

Cutaneotrichosporon dermatis

18.18

1.06

Cystobasidium lysinophilum

9.10

0.71

Cystobasidium slooffiae

18.18

1.06

Cystofilobasidium infirmominiatum

9.10

0.18

Filobasidium magnum

18.18

0.35

Filobasidium wieringae

9.10

0.18

Hannaella luteola

9.10

0.53

Meira nashicola

54.55

3.72

Moesziomyces aphidis

27.27

5.49

Naganishia adeliensis

9.10

0.18

Naganishia albida

45.45

1.59

Naganishia cerealis

9.10

0.18

Naganishia diffluens

18.18

0.35

Papiliotrema aspenensis

36.36

4.42

Papiliotrema flavescens

27.27

2.48

Rhodotorula graminis

90.91

12.04

Rhodotorula mucilaginosa

190

100.00

33.63

Rhodotorula toruloides

18.18

0.53

Ustilago filiformis

9.10

0.18

Ustilago longissima

9.10

0.18

Vishniacozyma carnescens

9.10

0.35

Vishniacozyma taibaiensis

9.10

0.35

Vishniacozyma victoriae

9.10

0.18

Total

565

CFU (units/L)

3 100

3 800

700

200

615

7 243

8 472

8 372

3 143

2 529

6 586

3 100

Sites	WA02	WA03	WA04	WA05	WA06	WA07	WA08	WA09	WA10	WA11	WA12
WA02	1
WA03	0.57	1
WA04	0.27	0.25	1
WA05	0.21	0.36	0.19	1
WA06	0.23	0.29	0.26	0.44	1
WA07	0.13	0.18	0.18	0.25	0.27	1
WA08	0.20	0.18	0.13	0.15	0.17	0.45	1
WA09	0.16	0.26	0.20	0.36	0.38	0.43	0.43	1
WA10	0.25	0.31	0.15	0.33	0.22	0.18	0.18	0.21	1
WA11	0.20	0.43	0.18	0.25	0.27	0.23	0.23	0.33	0.44	1
WA12	0.09	0.17	0.05	0.13	0.14	0.11	0.11	0.08	0.40	0.25	1

Sites

WA02

WA03

WA04

WA05

WA06

WA07

WA08

WA09

WA10

WA11

WA12

WA02

WA03

0.57

WA04

0.27

0.25

WA05

0.21

0.36

0.19

WA06

0.23

0.29

0.26

0.44

WA07

0.13

0.18

0.25

0.27

WA08

0.20

0.18

0.13

0.15

0.17

0.45

WA09

0.16

0.26

0.20

0.36

0.38

0.43

WA10

0.25

0.31

0.15

0.33

0.22

0.18

0.21

WA11

0.20

0.43

0.18

0.25

0.27

0.23

0.33

0.44

WA12

0.09

0.17

0.05

0.13

0.14

0.11

0.08

0.40

0.25

Sites	T/℃	Turb	pH	TP (mg/L)	TN (mg/L)	TOC (mg/L)	TH (mg/L)	EC (μS/cm)
The letters in the table indicate size and significant differences: “a” is the largest, “ab” is the second largest, and decreases in sequence from beginning to end; There is a significant difference between “a” “b” and “c”, with aP-value of < 0.05. “ab” indicates that there is no significant difference between “a” and “b”, while “bc” indicates that there is no significant difference between “b” and “c”, but there is a significant difference between “a” and “ab”, and so on.
WA02	11.0c	7.01bc	7.67c	0.06bc	0.99b	30.73c	189.00ab	343.00aab
WA03	15.0ab	6.51bc	7.90b	0.06bc	0.75bc	37.97bc	131.00b	257.00bc
WA04	16.0a	26.77ab	7.90b	0.09b	1.02ab	39.93ab	134.33ab	274.00b
WA05	16.0a	12.61ab	7.80bc	0.26a	2.06a	42.87ab	132.00ab	289.33ab
WA06	16.0a	34.07a	7.90b	0.14ab	1.29ab	40.63ab	136.00ab	283.00ab
WA07	16.0a	13.01ab	7.90b	0.11ab	0.97bc	41.07ab	124.33bc	279.33ab
WA08	16.0a	8.05b	8.00ab	0.07bc	0.71bc	38.07bc	127.00bc	277.00ab
WA09	16.0a	12.71ab	8.07ab	0.13ab	0.69bc	39.63bc	144.00ab	273.00bc
WA10	15.0ab	6.25bc	8.10a	0.10ab	0.56c	39.70b	117.00bc	267.33bc
WA11	15.0ab	4.30c	8.10a	0.06c	0.77bc	35.80bc	115.33c	251.00c
WA12	14.5b	21.90ab	7.90b	0.10ab	1.13ab	70.60a	228.00a	465.00a

Sites

T/℃

Turb

TP (mg/L)

TN (mg/L)

TOC (mg/L)

TH (mg/L)

EC (μS/cm)

The letters in the table indicate size and significant differences: “a” is the largest, “ab” is the second largest, and decreases in sequence from beginning to end; There is a significant difference between “a” “b” and “c”, with aP-value of < 0.05. “ab” indicates that there is no significant difference between “a” and “b”, while “bc” indicates that there is no significant difference between “b” and “c”, but there is a significant difference between “a” and “ab”, and so on.

WA02

11.0c

7.01bc

7.67c

0.06bc

0.99b

30.73c

189.00ab

343.00aab

WA03

15.0ab

6.51bc

7.90b

0.06bc

0.75bc

37.97bc

131.00b

257.00bc

WA04

16.0a

26.77ab

7.90b

0.09b

1.02ab

39.93ab

134.33ab

274.00b

WA05

16.0a

12.61ab

7.80bc

0.26a

2.06a

42.87ab

132.00ab

289.33ab

WA06

16.0a

34.07a

7.90b

0.14ab

1.29ab

40.63ab

136.00ab

283.00ab

WA07

16.0a

13.01ab

7.90b

0.11ab

0.97bc

41.07ab

124.33bc

279.33ab

WA08

16.0a

8.05b

8.00ab

0.07bc

0.71bc

38.07bc

127.00bc

277.00ab

WA09

16.0a

12.71ab

8.07ab

0.13ab

0.69bc

39.63bc

144.00ab

273.00bc

WA10

15.0ab

6.25bc

8.10a

0.10ab

0.56c

39.70b

117.00bc

267.33bc

WA11

15.0ab

4.30c

8.10a

0.06c

0.77bc

35.80bc

115.33c

251.00c

WA12

14.5b

21.90ab

7.90b

0.10ab

1.13ab

70.60a

228.00a

465.00a