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辽西北风沙区是典型的生态脆弱区,位于我国科尔沁沙地南缘,该区水资源不足,气候干燥,植被覆盖率低,大风频繁,是三北防护林体系建设工程的重点治理区域。自20世纪50年代以来,我国一直在实施大规模沙漠化治理工程[1],进行大面积人工造林[2]。1978年以来,我国启动了世界上最大的造林项目“三北防护林带发展计划”[3]。近四十年来,该地区营造了大面积防护林,在生态修复中起着重要的作用。防护林体系的构建是改善沙区生态环境的有效措施,不仅有助于减轻风沙危害[4],防止土壤退化,而且防护林种植有助于改良土壤,增加土壤有机质含量[5],从而影响土壤微生物的活性。
土壤微生物是陆地生态系统的重要生物驱动力之一[6],是土壤生物地球化学循环中最核心的部分[7],是控制生态系统中C、N和其他养分流的关键[8],在推动土壤生态系统的平衡稳定发展方面发挥着至关重要的作用[9]。而且,土壤微生物也是最敏感的指标之一[10-12],较土壤理化特性更能反映土壤质量。诸多研究表明,植被类型、土壤特性以及土壤微生物具有一定的相关性[13-14],土壤理化性质及养分含量的改变会显著影响微生物群落功能多样性[15-16],土壤微生物在提高土壤氮、磷、钾,改善土壤质量等方面也具有重要作用[17]。微生物多样性较高表明土壤养分补充性能较好,且微生物对底物的利用率高[18]。其中,土壤真菌是土壤中的主要分解者之一,能够分解土壤中的有机物(植物残体),为植物提供养分[19],部分真菌有助于提高植物的抗逆性,维持植物正常生长。同时,土壤真菌在促进土壤稳固、团聚体的形成以及改善土壤结构方面具有重要作用[20],对土壤环境乃至整个生态系统都会产生重要影响。在生态系统恢复过程中,真菌群落的变化是一个关键性指标[21],不同树种土壤真菌数量和群落结构具有显著差异[19,22]。除真菌群落多样性和结构之外,微生物功能也是反映土壤质量的重要因子。其中,FUNGuild软件的开发利用有助于我们更好地研究真菌功能类群,且已被众多的学者用于真菌功能群的研究[23-24]。
目前,针对辽西北风沙区防护林的研究主要集中在不同防护林土壤C、N、P垂直分布特征[25],土壤水稳性团聚体质量分数和有机质质量分数方面[26],而针对该地区不同防护林土壤真菌的多样性及功能尚无系统报道。因此,本研究以辽宁省西北部的昌图县付家林场典型的人工防护林,樟子松人工林(Pinus sylvestris var. mongolica Litv.)、油松人工林(P. tabuliformis Carrière)、杨树人工林(Populus ×canadensis Moench)为研究对象,探讨不同人工防护林下土壤真菌群落结构和功能特征,以期为该地区植被恢复和人工林固沙造林树种的选择提供参考。
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试验地位于辽宁铁岭市昌图县付家林场,地处科尔沁沙地东南边缘,位于辽宁、内蒙古、吉林三省区交汇处,地势平坦,地貌属辽河冲积平原兼有少量沙丘。土壤类型大部分为风沙土。该地属温带半湿润半干旱大陆性季风气候,最高气温为35.6 ℃,最低气温为−31.5 ℃,降雨集中在7—8月,年平均降水量400~550 mm,年蒸发量1 843 mm,远高于年降水量,极度干旱。该地区的原有植被为少量的灌木和草本,现植被以樟子松和杨树等形成的防风固沙林为主(表1)。
表 1 样地信息
Table 1. Sampling site information
树种
Tree species林分密度
Stand density/
(plant·hm−2)树高
Height/m胸径
Diameter at breast height/cm郁闭度
Crown density/%杨树人工林
Populus × canadensis1 100 15.10 13.30 60 樟子松人工林
Pinus sylvestris var. mongolica275 14.10 33.56 65 油松人工林
Pinus tabuliformis575 10.56 18.50 70 -
通过测定,发现昌图县付家林场不同类型人工林的土壤化学性质差异显著(见表2),杨树人工林土壤pH值为5.92,显著高于油松和樟子松人工林(P<0.05);杨树人工林土壤可溶性有机碳和速效磷的含量分别为105.46 g·kg−1和16.00 g·kg−1,均显著高于樟子松和油松人工林(P<0.05),樟子松人工林下土壤的全氮含量最高,为1.05 g·kg−1,显著高于杨树和油松人工林(P<0.05),杨树人工林土壤C/N为9.15,显著低于樟子松和油松人工林(P<0.01)。通过比较,发现杨树人工林能显著增加土壤pH、土壤可溶性有机碳和速效磷的含量,降低土壤C/N。
表 2 不同人工林土壤特性
Table 2. Soil properties for different plantation forests
树种
Tree speciespH值
pH value可溶性有机碳
DOC/(g·kg−1)全氮
Total N/(g·kg−1)碳氮比
C/N ratio速效磷
Available P/(mg·kg−1)杨树人工林
Populus ×canadensis5.92±0.11aA 105.46±7.81aA 0.85±0.08bA 9.15±0.26bB 16.00±1.80aA 樟子松人工林
Pinus sylvestris var. mongolica5.57±0.21bA 83.42±18.42abA 1.05±0.10aA 11.41±0.31aA 3.77±1.43bB 油松人工林
Pinus tabuliformis5.53±0.17bA 80.10±32.32bB 0.97±0.07abA 11.79±0.49aA 3.09±0.98bB 注:表中数据为均值±标准差(n=3),不同小写字母表示不同处理之间差异显著(P<0.05),不同大写字母表示不同处理之间差异显著(P<0.01)。 -
通过对9个样地的土壤样品DNA测序分析,以97%相似性阀值进行OTU聚类,获得真菌群落结构组成信息。真菌测序深度(Coverage)为97%,与数据库进行比对后可知,真菌包括15个门、54个纲和513个属。测序结果经去除嵌合体和低质量的序列后,从所有样品中共得到真菌ITS序列494,675条,平均每个土壤样品有54 963条序列,平均长度为330 bp。为了确定样品的稀疏曲线,从每个样本中随机选择21 490个reads,在3%差异水平下,随着实测序列数量的增加,曲线趋于平坦,表明本试验获得了大部分样本信息,能够反映不同人工林土壤真菌群落组成(图1)。当序列相似度为97%时,在真菌门水平上,樟子松人工林、油松人工林和杨树人工林分别有579、464和708个OTU(表3)。基于Unweighted UniFrac距离的NMDS分析结果,发现樟子松人工林和油松人工林土壤真菌群落结构具有较大的相似性,都位于NMDS1轴的负半轴,而杨树人工林真菌群落位于NMDS1轴的正半轴,与樟子松和油松人工林土壤真菌群落明显沿NMDS1轴分开(图2)。
图 1 3个真菌群落高通量测序的ITS rDNA稀疏曲线
Figure 1. Rarefaction curves of ITS rDNA for high throughput sequencing of three fungal communities
图 2 不同人工林土壤真菌群落非度量多维尺度分析
Figure 2. Non-metric multidimensional scaling (NMDS) analysis based on Unweighted UniFrac distance under different plantation forests
表 3 不同人工林土壤真菌多样性指数
Table 3. Soil fungal diversity index in different plantation forests
树种
Tree speciesOTUs Simpson指数
Simpson indexChao 1指数
Chao1 indexACE指数
ACE indexShannon指数
Shannon index樟子松人工林
Pinus sylvestris var. mongolica579 0.962±0.018abAB 617.97±40.91bB 622.58±49.00bB 6.60±0.66aA 油松人工林
Pinus tabuliformis464 0.942±0.004bB 472.46±57.46cC 471.26±56.13cC 6.14±0.51aA 杨树人工林
Populus ×canadensis708 0.980±0.003aA 905.96±33.74aA 931.47±38.63aA 6.73±0.77aA 注:表中数据为均值±标准差(n=3),不同小写字母表示不同处理之间差异显著(P<0.05),不同大写字母表示不同处理之间差异显著(P<0.01)。 通过分析发现,付家林场不同人工林土壤真菌群落多样性指数存在显著差异,杨树人工林土壤真菌的Simpson指数、Chao1指数和ACE指数分别为0.980、905.96和931.47,都显著高于其他(P < 0.05);油松人工林最低,分别为0.942、472.46和471.26。不同人工林土壤真菌的Shannon指数无显著差异(表3)。土壤pH值与ACE指数(r = 0.81,P < 0.01)和Chao1指数(r = 0.81,P < 0.01)呈极显著正相关;土壤DOC与土壤真菌Simpson指数(r = 0.86,P < 0.01),ACE指数(r = 0.79,P < 0.05)和Chao1指数(r = 0.78,P < 0.05)呈显著正相关。土壤AP与土壤真菌ACE指数(r = 0.86,P < 0.01)和Chao1指数(r = 0.86,P < 0.01)呈极显著正相关,土壤C/N与土壤真菌Simpson指数(r = −0.80,P<0.01),ACE(r = −0.91,P<0.01)和Chao1指数(r = −0.91,P < 0.01)呈极显著负相关(图3)。
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研究发现,付家林场不同样地土壤真菌群落组成存在差异,根据真菌的高级分类可知[28],所得序列属于14门(不包括unidentified),优势菌门为子囊菌门(Ascomycota)和担子菌门(Basidiomycota),它们的相对丰度分别为49.58% ~ 59.56%和23.99% ~ 29.77%。其他相对丰度占1%以上的门分别为被孢霉门(Mortierellomycota) (3.89%)、油壶菌门(Olpidiomycota)(1.28%)和壶菌门(Chytridiomycota)(1.21%)。油松人工林中子囊菌门的相对丰度最高,而油松人工林土壤中担子菌门、被孢霉门和壶菌门的相对丰度均最低(图4)。
图 4 不同人工林土壤优势真菌门的相对丰度
Figure 4. The relative abundances of dominant fungal communities at phylum level
在纲水平上,平均相对丰度大于1%的纲共有9个,分别为座囊菌纲(Dothideomycetes)(13.86%~20.68%)、伞菌纲(Agaricomycetes)(12.83%~22.28%)、粪壳菌纲(Sordariomycetes) (6.38%~23.09%)、散囊菌纲(Eurotiomycetes) (0.26%~23.80%)、银耳纲(Tremellomycetes)(5.93%~13.55%)、锤舌菌纲(Leotiomycetes) (5.44% ~6.68%)、被孢霉纲(Mortierellomycetes) (2.13%~4.83%)、盘菌纲(Pezizomycetes)(0.22%~7.06%)和GS18 (0.17% ~2.58%)。其中,座囊菌纲、伞菌纲、锤舌菌纲、被孢霉纲和盘菌纲的相对丰度在不同人工林样地中无显著差异。在杨树人工林土壤中粪壳菌纲和银耳纲的相对丰度最高,且显著高于其他人工林样地(P<0.05),然而,在油松人工林土壤中散囊菌纲相对丰度含量最高,显著高于其他样地(P<0.05) (图5)。
在所有的样品中共检测出513个属,其中,相对丰度大于1%的主要的真菌类群为Guehomyces (3.96%)、被孢霉属(Mortierella) (3.88%)、青霉菌属(Penicillium) (3.31%)、Plectosphaerella (2.52%)、小皮伞属(Marasmius) (2.50%)、Knufia (2.33%)、Phallus (2.31%)、Devriesia (1.54%)、红菇属(Russula) (1.48%),Trechispora (1.37%)、Microidium (1.35%)、Lectera (1.17%)和篮状菌属(Talaromyces)(1.10%)(图6)。相对丰度前50的真菌属的热图分析表明,不同植被类型的真菌群落结构不同,可以划分为两个类群,其中樟子松人工林和油松人工林为一类,杨树人工林为单独一类(图7)。
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真菌的生态功能类群是基于营养型来划分的。根据FUNGuild数据库的比对结果,对3种人工林真菌的营养型(trophic mode)进行分类统计,结果如图3所示,检测到8个营养型(Trophic mode),分别为致病菌-腐生-共生营养型(Pathogen-Saprotroph-Symbiotroph)、病理营养型(Pathotroph)、病理-腐生营养型(Pathotroph-Saprotroph)、病理-腐生-共生营养型(Pathotroph-Saprotroph-Symbiotroph)、病理-共生营养型(Pathotroph-Symbiotroph)、腐生营养型(Saprotroph)、腐生-共生营养型(Saprotroph-Symbiotroph)和共生营养型(Symbiotroph)。该区3种人工林土壤真菌主要以腐生营养型为主,腐生-共生营养型次之。对不同人工林土壤真菌功能类群进行聚类分析结果表明,不同人工林真菌群落功能可划分为两个类群,其中油松人工林和红松人工林土壤真菌群落功能为一类,杨树人工林土壤真菌功能类群为一类(图8)。
图 8 不同人工林土壤真菌群落功能热图
Figure 8. The heatmap of the fungal community functions in different plantation forest soils.
为研究土壤真菌群落功能与土壤环境因子间的相关性,采用CCA分析对土壤环境因子和真菌群落功能类的关系进行限制性排序分析,如图9所示,土壤pH (r=0.76)、可溶性有机碳(r=0.61)、C/N (r=−0.83)和速效磷(r=−0.75)与CCA1轴的相关性较大,第一轴的解释量为53.0%。土壤速效磷(r=−0.54)与第二轴的相关性较大,解释量为36.5%。由此可见,土壤pH、可溶性有机碳、C/N和速效磷为土壤真菌群落功能的主要影响因子。
辽西北风沙区不同人工林土壤真菌群落结构及功能特征
Studies on Soil Fungal Community Composition and Function Characteristics of Different Plantations of Sandy Area, Northwest Liaoning Province
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摘要:
目的 为了揭示土壤真菌群落结构及功能对不同人工林的响应特征和响应机制,评估不同人工林对地下生态系统的影响。 方法 以辽西北风沙区樟子松人工林、油松人工林和杨树人工林为研究对象,对每个样地采用“S”型布点采样,采集0~10 cm表层土壤并进行混合,采用Illumina Miseq高通量测序技术和FUNGuild软件,获取并分析辽西北风沙区不同人工林土壤真菌群落组成与功能特征,并对土壤真菌群落多样性与土壤环境因子进行相关性分析。 结果 表明:杨树人工林能显著增加土壤pH值、土壤可溶性有机碳和速效磷的含量,降低土壤的C/N。该区人工林土壤优势真菌门类群是子囊菌门和担子菌门,优势真菌属类群为Guehomyces、被孢霉属及青霉菌属。NMDS和heatmap分析结果表明,不同人工林土壤真菌群落结构和功能特征差异显著,都可划分为两个聚类,其中,樟子松人工林和油松人工林土壤真菌群落结构和功能的相似性较大,而杨树人工林土壤真菌群落与针叶林土壤真菌群落结构和功能的差异较大。土壤pH、速效磷、DOC和C/N是影响土壤真菌多样性和土壤真菌群落功能的主要因子。 结论 不同人工林对土壤特性及土壤真菌群落的结构和功能特征具有不同的影响,杨树人工林显著提高土壤养分有效性,增加土壤真菌多样性。因此,在辽西北风沙区种植杨树较针叶树能更好地改善土壤。 Abstract:Objective To reveal the response characteristics and mechanism of soil fungal community structure and function to different plantations, and to evaluate the impact of different plantations on the underground ecosystem. Method Pinus sylvestris var. mongolica plantation, Pinus tabuliformis plantation, and Populus × canadensis plantation in the north wind sand area of western Liaoning province were investigated. For each sampling plot, the "S" type spot sampling method was used to collect the surface soil of 0 ~ 10 cm at multiple points.Illumina Miseq high-throughput sequencing technology and FUNGuild software were applied to obtain and analyze the response characteristics of soil microbial community composition and function in different plantations. Meanwhile, the relationships between soil fungal community diversity, community structure and soil environmental factors were analyzed. Result The plantation of Populus × canadensis could significantly increase the soil pH value, soil-dissolved organic carbon and available phosphorus content, and reduce soil C/N ratio. The dominant fungal phyla in these plantations were Ascomycota and Basidiomycota, and the dominant fungal genera were Guehomyces, Mortierella and Penicillium. The results of NMDS and heatmap analysis showed that there were significant differences in the community structure and function of soil fungi among different plantations and the community composition and function could be divided into two clusters. The community structure of soil fungi in Pinus sylvestris var. mongolica, and Pinus tabuliformis plantations were similar, while the community composition and function in plantation of Populus × canadensis Moench were significantly different from that in coniferous plantations. The soil pH, available phosphorus, dissolved organic carbon and C/N ratio were the main factors affecting soil fungal community diversity and functions. Conclusion The effect on soil characteristics and the soil fungal composition and function differs among different plantations. The plantation of Populus × canadensis Moench significantly increases the soil nutrient availability and soil fungal diversity. -
表 1 样地信息
Table 1. Sampling site information
树种
Tree species林分密度
Stand density/
(plant·hm−2)树高
Height/m胸径
Diameter at breast height/cm郁闭度
Crown density/%杨树人工林
Populus × canadensis1 100 15.10 13.30 60 樟子松人工林
Pinus sylvestris var. mongolica275 14.10 33.56 65 油松人工林
Pinus tabuliformis575 10.56 18.50 70 表 2 不同人工林土壤特性
Table 2. Soil properties for different plantation forests
树种
Tree speciespH值
pH value可溶性有机碳
DOC/(g·kg−1)全氮
Total N/(g·kg−1)碳氮比
C/N ratio速效磷
Available P/(mg·kg−1)杨树人工林
Populus ×canadensis5.92±0.11aA 105.46±7.81aA 0.85±0.08bA 9.15±0.26bB 16.00±1.80aA 樟子松人工林
Pinus sylvestris var. mongolica5.57±0.21bA 83.42±18.42abA 1.05±0.10aA 11.41±0.31aA 3.77±1.43bB 油松人工林
Pinus tabuliformis5.53±0.17bA 80.10±32.32bB 0.97±0.07abA 11.79±0.49aA 3.09±0.98bB 注:表中数据为均值±标准差(n=3),不同小写字母表示不同处理之间差异显著(P<0.05),不同大写字母表示不同处理之间差异显著(P<0.01)。 表 3 不同人工林土壤真菌多样性指数
Table 3. Soil fungal diversity index in different plantation forests
树种
Tree speciesOTUs Simpson指数
Simpson indexChao 1指数
Chao1 indexACE指数
ACE indexShannon指数
Shannon index樟子松人工林
Pinus sylvestris var. mongolica579 0.962±0.018abAB 617.97±40.91bB 622.58±49.00bB 6.60±0.66aA 油松人工林
Pinus tabuliformis464 0.942±0.004bB 472.46±57.46cC 471.26±56.13cC 6.14±0.51aA 杨树人工林
Populus ×canadensis708 0.980±0.003aA 905.96±33.74aA 931.47±38.63aA 6.73±0.77aA 注:表中数据为均值±标准差(n=3),不同小写字母表示不同处理之间差异显著(P<0.05),不同大写字母表示不同处理之间差异显著(P<0.01)。 -
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