刘茗(1992—), 女, 安徽宿州人, 硕士研究生, 主要从事土壤生物与生物化学研究。E-mail:
为研究亚热带不同森林植被类型土壤固碳微生物特征及其影响因子,选取毛竹林(Moso banboo groves)、阔叶林(Broad-leaved forest)、杉木林(Chinese fir forest)和马尾松林(Masson pine forest)等4种森林植被为研究对象,以
To alleviate the global climate change is one of the most important environmental challenges facing mankind. Autotrophic microorganisms, especially those in forest ecosystems, have been reported to have a strong ability to adapt to environmental changes and a high potential to sequestrate carbon. However, how their carbon-sequestrating effects vary with type of forests vegetation remains unclear. To explore this complex relationship, investigations were made of populations and community structures of soil carbon-sequestrating bacteria in the soils of four typical subtropical forests.
Soil samples were collected from the surface (0-20 cm) and subsurface (20-40 cm) soil layers in the four types of forests, that is Moso banboo, Broad-leaved trees, Chinese fir and Masson pine for analysis of
Abundance of the bacteria with 16S rRNA genes and those with
To sum up, all the findings in this study indicate that population and community structure diversity of the soil carbonsequestrating microbes varied with type of the vegetation. Comparison of the four forest soils in soil physico-chemical property, and genetic abundance, diversity and community structure of the carbon-sequestrating bacteria shows Moso bamboo groves are the best type of vegetation benefitting soil fertility and carbon-sequestrating bacteria. Contribution of the bacteria to accumulation of organic matter was higher in the soil under bamboo groves than under broad-leaved forests, but as to how much, further investigations should be done.
工业飞速发展以及人类活动加剧了全球性温室效应[
综上所述,从整个生物圈的物质、能量流动来看,研究自养微生物的固碳功能具有重要意义,但影响固碳微生物因素却较为复杂。森林是陆地重要的碳汇,全球森林生态系统存储碳量高达650 Gt,其中地下土壤占45%[
本研究以植物多样性丰富的亚热带不同森林植被类型为对象,选取亚热带典型森林植被毛竹林(Moso banboo groves)、阔叶林(Broad-leaved forest)、杉木林(Chinese fir forest
研究样地位于浙江省杭州市临安区玲珑山(30°14′N,119°42′E),属中亚热带季风气候,热量丰富,雨量丰沛,光照充足,四季分明,具有春多雨、夏湿热、秋气爽、冬干冷的气候特征。多年平均降雨量1 628.6 mm,降水日158 d,全年日照时数1 847 h,无霜期237 d,全年平均气温16.4 ℃。土壤为发育于凝灰岩的黄红壤,具有典型的中亚热带的森林生态系统和森林景观,植被主要包括毛竹、阔叶、杉木、马尾松等典型亚热带植被。其中,阔叶林、杉木林和马尾松林下植被均包含丛生灌木,有蕨类植物,毛竹林下植被很少,仅有稀疏的草本植物。
于2017年10月按照生态控制原则,选择同一成土母质的毛竹林(MB)、阔叶林(BL)、杉木林(CF)和马尾松林(MP)。分别于各个林分的下坡位、中坡位、上坡位确定采样区,在采样区周围10 m2范围内按照三角形确定3个取样点,分别采集表层(0~20 cm)及亚表层(20~40 cm)土壤混匀得到一个土样,每种林分选择4个重复,4种林分共得32个(4×4×2)土样。新鲜土样去除植物残体和大的石头过筛(2 mm)后装入密封袋,将样品放入冰盒带回实验室。样品共分为2份,1份测定含水率后鲜土提取土壤细菌总DNA,并冷冻干燥存放于-70 ℃冰箱,用于其他分子生物学分析;另一份于室内自然风干,充分研磨过筛后用于土壤基本理化性质的测定。
土壤理化性质分析:分析方法参考文献[
土壤总DNA提取:采用Power Soil DNA Isolation Kit试剂盒提取土壤总DNA。测定鲜土的含水率后,换算成0.35 g干土重,称取相应的新鲜土样,按试剂盒说明书进行提取。DNA提取成功后经1%(m/v)的琼脂糖凝胶电泳检测DNA片段大小,并用微量分光光度计检测其浓度和纯度。提取后的DNA样品保存于-40 ℃冰箱。
实时荧光定量PCR(Real-time Quantutative PCR,qPCR),测定固碳功能菌
高通量测序:采用同上K2f/V2r引物对,对土壤DNA进行扩增,将PCR产物纯化后送至上海派森诺公司,采用Illumina MiSeq平台对群落DNA片段进行双端(Paired-end)测序。下机数据选择NCBI(National Center for Biotechnology Information)数据库的氨基酸序列进行对比,进行物种注释得到分类鉴定结果。
利用Microsoft Excel 2010软件对数据进行初处理,采用SPSS 21.0软件对数据进行和双因素方差分析(Two-Way ANOVA)和最小显著差数(LSD)法进行显著性和多重比较(α=0.05),采用皮尔逊(Pearson)进行相关性分析。根据OTU表计算α多样性指数。利用Origin软件绘制基因丰度图和群落结构丰度图。使用Canoco 5.0软件对固碳细菌群落结构与环境因子的相关性进行冗余分析(Redundancy analysis,RDA)。
4种林分虽然发育于同一母岩,但土壤理化性质却存在差异(
不同林分土壤理化性质(均值±标准差)
Soil physico-chemical propert ties relative to type of forest(mean ± SD)
土层 |
林分 |
pH | 有机碳 |
全氮 |
碳氮比 |
碱解氮 |
有效磷 |
速效钾 |
注:MB:毛竹,BL:阔叶林,CF:杉木林,MP:马尾松林;*, | ||||||||
0~20 | MB | 4.77±0.15aA | 17.68±2.87aA | 1.55±0.17aA | 11.57±1.87aA | 34.0±4.42abA | 6.47±1.31aA | 102±22.0aA |
BL | 4.38±0.06cA | 23.90±4.21aA | 1.95±0.37aA | 12.16±1.59aA | 40.6±4.39aA | 6.05±0.72aA | 97.0±12.2aA | |
CF | 4.60±0.14abA | 14.43±2.23bA | 1.30±0.14bA | 11.19±2.20aA | 27.0±7.68bcA | 2.15±0.98bA | 76.0±5.35aA | |
MP | 4.42±0.11bcA | 15.05±2.86bA | 1.25±0.13bA | 12.23±2.48aA | 25.4±1.75cA | 4.45±2.34aA | 82.0±20.3aA | |
20~40 | MB | 4.74±0.09aA | 13.48±2.82aA | 1.48±0.22aA | 9.12±0.81aA | 25.7±4.90aA | 4.71±0.65bB | 76.3±12.8aA |
BL | 4.42±0.10bA | 12.60±3.60aB | 1.20±0.36aA | 11.3±4.32aA | 25.9±3.86aB | 4.13±0.80abB | 83.3±8.26aA | |
CF | 4.67±0.06aA | 11.93±5.49aA | 1.05±0.37aA | 10.6±1.42aA | 23.2±2.17aA | 1.37±0.66cB | 74.5±7.72aA | |
MP | 4.37±0.13bA | 12.93±2.45aA | 1.18±0.29aA | 11.0±1.57aA | 20.7±2.58aB | 3.12±1.28bA | 83.5±13.5aA | |
F值F value | ||||||||
林分Forest stand | 21.3*** | 3.33* | 4.46* | 0.64 | 8.99*** | 1.97 | 15.9** | |
土层Layer | 0.05 | 16.9*** | 8.84** | 2.60 | 26.3*** | 4.08 | 11.4** | |
交互Interaction | 0.59 | 3.04* | 2.72 | 0.27 | 2.62 | 1.63 | 0.35 |
采用荧光实时定量PCR分析,4种林分土壤细菌16S rRNA基因和固碳细菌
不同林分土壤细菌16S rRNA(A)、固碳细菌
Abundance of 16S rRNA(A)and
为揭示土壤理化性质对不同林分土壤细菌16S rRNA和固碳细菌
土壤细菌16S rRNA和固碳细菌
Pearson's correlations(r)of 16S rRNA and
林分 |
功能基因 |
pH | SOC | TN | C:N | AN | AP | AK |
注:以林分为单位分析时, | ||||||||
MB | 16S rRNA | 0.673 | 0.142 | -0.047 | 0.269 | 0.394 | 0.498 | 0.423 |
0.391 | 0.538 | 0.448 | 0.407 | 0.427 | 0.373 | 0.032 | ||
BL | 16S rRNA | -0.406 | 0.333 | 0.161 | 0.250 | 0.462 | 0.826* | 0.017 |
-0.253 | 0.154 | -0.183 | 0.479 | 0.315 | 0.767* | -0.291 | ||
CF | 16S rRNA | 0.252 | 0.123 | 0.209 | -0.042 | -0.236 | -0.323 | 0.042 |
0.614 | -0.272 | -0.054 | -0.426 | -0.484 | 0.022 | -0.395 | ||
MP | 16S rRNA | 0.167 | -0.328 | -0.656 | 0.332 | -0.064 | 0.0439 | 0.519 |
0.768* | -0.098 | -0.524 | 0.539 | 0.181 | 0.587 | 0.744* | ||
土层Layer/cm | 功能基因Functional gene | pH | SOC | TN | C:N | AN | AP | AK |
0~20 | 16S rRNA | 0.720** | -0.227 | -0.173 | -0.098 | -0.028 | 0.362 | 0.439 |
0.683** | -0.221 | -0.256 | 0.073 | -0.158 | 0.368 | 0.394 | ||
20~40 | 16S rRNA | 0.525* | 0.126 | 0.281 | -0.146 | 0.224 | -0.003 | 0.444 |
0.514* | 0.196 | 0.369 | -0.172 | 0.321 | 0.210 | 0.378 |
对4种林分土壤固碳细菌进行高通量测序,毛竹林、阔叶林、杉木林和马尾松林表层土壤分别测得36 598、33 599、35 093和35 129条序列,亚表层土壤测得32 552、33 783、30 381和38 502条有效序列,将测序结果在97%相似度下进行聚类得到OTU的代表序列,并与NCBI数据库比对进行物种注释,置信度阈值设为0.8,认为测序深度已经基本覆盖到样品中所有的物种。经比对鉴定得到4个门、6个纲、12个目、20个科、35个属、40个种的土壤固碳细菌群落信息。
基于OTU的α多样性指数可反应土壤固碳细菌群落结构的多样性。从表层土壤固碳细菌α多样性4个指标看(
不同林分土壤固碳细菌
土层 |
林分 |
辛普森Simpson | Chao1 | ACE | 香农Shannon |
0~20 | MB | 0.97±0.01 aB | 1 345 ±171 aA | 13 64±140 aA | 8.11±0.10 aA |
BL | 0.97±0.00 aA | 1 366±10.2 aB | 1 472±17.0 aB | 8.03±0.23 aA | |
CF | 0.63±0.07 bB | 872.4±268 bB | 961.2±294 bB | 4.21±0.88 bB | |
MP | 0.98±0.02 aA | 1 302±129 aA | 1 379±168 aA | 8.47±0.54 aB | |
20~40 | MB | 0.99±0.00 aA | 1 638±305 aA | 1 737±323aA | 8.73±0.54 aA |
BL | 0.96±0.01 aA | 1 522±60.2 aA | 1 613±24.5 aA | 7.94±0.11 abA | |
CF | 0.89±0.10 aA | 1 443±323 aA | 1 555±353 aA | 7.13±1.51 bA | |
MP | 0.99±0.00 aA | 1 782±421 aA | 1 965±503 aA | 9.04±0.34 aA | |
F值F value | |||||
林分Forest stand | 51.4*** | 5.27** | 4.54* | 39.7*** | |
土层Layer | 26.5*** | 27.9*** | 28.5*** | 25.8*** | |
交互Interaction | 23.6*** | 1.83 | 1.88 | 11.8*** |
为分析4种不同林分土壤固碳细菌群落组成的差异,以固碳细菌OTU种类数为依据,对其分组构建韦恩图分析不同处理之间的“相似关系”(
不同林分土壤固碳细菌OTUs韦恩图
OTUs Veen of soil carbon sequestrating bacteria relative to forest stand
在属分类水平上(
不同林分土壤固碳细菌在属水平上的群落结构组成分布
Composition of the soil carbon sequestrating bacteria community structure at genus level relative to forest stand
为分析4种林分土壤理化因子对土壤固碳细菌群落结构的影响,选取表层及亚表层土壤具有代表性的优势菌属(聚类大于1%)为物种变量、土壤理化性质为环境变量进行冗余分析,根据蒙特卡洛检验,选取影响较大的环境因子分析。结果表明,表层土壤第一排序轴和第二排序轴分别解释了34.4%和20.7%的变异(
土壤固碳细菌在属水平上与环境因子的冗余分析
Redundancy analysis of soil carbon sequestrating bacteria at the genus level with environmental parameters relative to forest stand
根据蒙特卡洛检验,选取亚表层土壤中(
通过qPCR分析亚热带典型森林植被土壤微生物16S rRNA基因与
不同生态系统因为土壤环境因子的巨大差异导致土壤固碳细菌丰度差异[
对固碳细菌功能基因
不同林分之间固碳细菌丰度和多样性差异的根本原因是群落结构不同,从韦恩图结果显示的林分之间差异(
本研究中4种林分表层土壤占绝对优势的甲基化石油杆菌属,在桂林毛村岩溶的岩溶区农田土壤中为第二优势属[
毛竹林、阔叶林、杉木林和马尾松林4种森林植被土壤中均存在相当数量的细菌及固碳细菌,其中毛竹林显著高于其他3种林分,阔叶林土壤有效磷以及不同土层的pH分别与对应的细菌与固碳细菌基因丰度显著相关。杉木土壤固碳细菌多样性显著低于其他3种林分;双因子分析显示,林型、土层之间均存在显著差异。4种林分土壤固碳细菌群落结构与农田、草地和海洋等生态系统有较大差异,甲基化石油杆菌属和诺卡菌属为4种林分共同的优势菌属;毛竹和杉木两种人工林土壤固碳细菌群落结构组成较为相似。RDA结果显示,不同林分土壤pH、有机碳、有效磷、全氮差异是驱动土壤固碳细菌群落特征形成的主要因素。毛竹林土壤肥力最好,固碳细菌丰度最高,不同林分之间土壤微生物对土壤有机质的积累的贡献存在差异,确切定量结论有待进一步研究。
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