李昌骏(2001—),男,四川邻水人,硕士研究生,主要从事土壤肥力研究。E-mail:
为探究生物质炭负载解钾菌对土壤微生物特性的影响,基于5个处理即空白(CK)、施用化学钾肥(KCl)、接种解钾菌(KSB)、施用生物质炭(BC)、施用生物质炭负载解钾菌(BC-KSB)的黑麦草盆栽耗竭试验,分析不同处理下土壤酶和微生物群落结构的响应特征。结果表明,BC-KSB相比其余施肥处理更有利于提高土壤脲酶、蔗糖酶、酸性磷酸酶和过氧化氢酶的活性,同时也提高了土壤细菌的物种多样性与菌群丰富度,并提高了土壤有益菌群(绿弯菌门、放线菌门、芽孢杆菌属和慢生根瘤菌属)的丰度,抑制了土壤致病菌群(变形菌门和罗河杆菌属)的繁殖。各施肥处理相比CK均显著提升了黑麦草干物质量,且以BC-KSB处理对黑麦草干物质量的提升最为显著。与CK和KCl相比,BC-KSB能显著提高土壤微生物生物量碳、微生物生物量氮、有机质、全氮和速效钾的含量(
Biochar loaded bacteria is a practical application of microbial immobilization. Biochar is considered as the ideal carrier of microorganisms in recent years because of its large specific surface area, pore diameter and good absorbability. Biochar loaded with
Based on pot experiment with ryegrass and five treatments: control (CK), chemical potassium fertilizer (KCl), inoculation of potassium solubilizing bacteria (KSB), application of biochar (BC) and potassium solubilizing bacteria loaded biochar (BC-KSB), it was analyzed that the change characteristics of soil urease, acid phosphatase, invertase and catalase enzyme activities and microbial community structure, also the effects of biochar-loaded potassium solubilizing bacteria composite materials on soil nutrients, soil pH, microbial biomass carbon, microbial biomass nitrogen and ryegrass yield were discussed.
The soil urease and acid phosphatase activity increased first and then decreased while the soil invertase and catalase activity decreased with an increase in the number of harvests for treatments with microbial agents or biochar. Compared with other fertilization treatments, BC-KSB was more conducive to the increase of soil urease, invertase, acid phosphatase and catalase activities. In terms of soil bacterial community, BC-KSB improved the species diversity and flora uniformity of soil bacteria, increased the abundance of soil beneficial bacteria (Chloroflexi, Actinobacteriota,
KSB loaded biochar has a positive impact on ryegrass yield, soil nutrients, soil enzyme activity, and bacterial community structure, which provides profound significance for fertilizing soil and improving soil ecological environment.
钾是植物生长必需的营养元素,其丰缺状况会对作物产量造成直接影响[
土壤酶是一种生物催化剂,是土壤肥力和土壤养分循环的重要表征指标[
基于此,本研究基于黑麦草盆栽耗竭试验,提出如下科学假设:(1)生物质炭负载解钾菌对土壤酶活性会有明显的促进作用;(2)生物质炭负载解钾菌会对土壤细菌群落和土壤养分产生积极的影响。针对上述科学假设,拟采用高通量测序技术探究在土壤钾素逐渐匮乏过程中二者施用对土壤细菌群落的影响,进而通过冗余分析探究土壤酶活性、黑麦草产量、土壤化学和生物性质与土壤细菌群落的相关性,以期为功能性微生物与生物质炭的维护和改善土壤生态系统功能提供理论依据。
供试土壤:2020年3月于雅安市老板山农田区域(29°58'12"N,102°58'34"E)采集酸性紫色土,其基本理化性质与魏巍等[
供试微生物:由茶园根际土壤分离纯化,经生理生化特性检测,16S rRNA序列分析比对,确定该菌种为微嗜酸寡养单孢菌(
供试黑麦草:品种为进口多年生黑麦草(
供试生物质炭:采购自浙江长三角聚农科技开发有限公司,采用玉米秸秆在厌氧环境以450~ 50℃,烧制2 h。其pH为9.58,全磷6.43 g·kg–1,全钾8.38 g·kg–1,碱解氮60.3 mg·kg–1,有效磷130.2 mg·kg–1,速效钾326.5 mg·kg–1;生物质炭表面元素组成采用飞纳台式扫描电镜能谱一体机(Phenom-world B.V./Phenom Prox,荷兰)测定,其元素组成为C 76.72%、O 15.78%、N 4.73%、P 2.5%、K 0.05%、S 0.22%。
供试肥料:尿素(N 46.2%)、过磷酸钙(P2O5 12%)和氯化钾(K2O 60%)。
生物质炭负载解钾菌复合材料:生物质炭与LB液体培养基以1︰100(w/v)的比例混合后,将微嗜酸寡养单孢菌(
本研究设空白(CK)、施用化学钾肥(KCl)、接种解钾菌(KSB)、施用生物质炭(BC)、施用生物质炭负载解钾菌(BC-KSB)5个处理。CK处理为不施钾肥的空白对照,KCl处理施加0.4 g氯化钾,KSB处理施加20 mL的解钾菌液(1×109 cfu·mL–1),BC处理施加20 g生物质炭,BC-KSB处理施加20 g生物质炭负载解钾菌复合材料(1×109 cfu·g–1),除KSB处理外,其余四个处理均施加20 mL灭菌后的解钾菌液,每个处理三次重复。采用塑料桶(直径15 cm,高度15 cm)种植黑麦草,每桶装2 kg风干土,连续种植5茬,每茬在种植前均会施加基准计量的氮磷肥(N 0.40 g·pot–1、P2O5 0.3 g·pot–1)。生长期间定期浇无菌水,保持土壤水分在田间持水量的75%左右,大棚培养温度保持在10~30℃之间。在每茬种植30 d后进行收获,测定黑麦草干物质量,并采集15 g盆栽土壤,在手动移除土壤中的植物碎屑和残根后进行土壤酶活性的测定。第五茬黑麦草收获后的土样不仅测定其土壤酶活性,同时还测定土壤基本理化性质,另有部分储存于4℃冰箱送往上海凌恩生物科技有限公司进行高通量测序。
每茬收集的土样采用靛酚蓝比色法、磷酸苯二钠比色法、3,5-二硝基水杨酸比色法和高锰酸钾滴定法分别测定其脲酶、酸性磷酸酶、蔗糖酶和过氧化氢酶活性[
使用E.Z.N.A.® Soil DNA Kit(Omega Bio-tek,Norcross,GA,美国)试剂盒提取样本微生物总DNA,1%琼脂糖凝胶电泳检测基因组DNA。细菌群落结构采用16S rRNA全长通用引物27F 5’-AGRGTTYGATYMTGGCTCAG-3’和1492R5’-RGY TACCTTGTTACGACTT-3’进行PCR扩增[
原始数据已上传至美国国家生物技术信息中心(NCBI)数据库(SRA accession number:PRJNA876764)。使用SMRT Link Analysis软件6.0版处理PacBio原始reads,以获得环形一致性序列(CCS)reads:参数设置最小通过次数为3,最小预测准确度为0.99。原始reads通过SMRT Portal处理,以筛选序列的长度(< 800或 > 2 500 bp)和质量。通过去除barcode、引物序列、嵌合体和包含10个连续的相同碱基的序列进行进一步过滤。使用UPARSE将分类操作单元(OTU)以98.65%的相似性阈值聚类,并使用UCHIME去除嵌合序列。RDP Classifier对Silva Release138.1 16S rRNA数据库使用70%的置信度阈值分析每个16S rRNA基因序列的系统发育关系。进行物种组成分析和α多样性分析前,以最低的OTU序列数组别为标准进行抽平处理。
基于高通量测序的OTU数据,使用Mothur v1.35.1计算细菌α多样性指数,利用香农(Shannon)指数和辛普森(Simpson)指数表征细菌的群落多样性,香农指数越高,微生物多样性越高;辛普森指数越高,微生物多样性越低;利用Chao1指数和多度覆盖的物种估计量(abundance-based coverage estimator,ACE)指数表征细菌的群落丰富度,Chao1指数值越高代表群落物种越丰富,ACE指数是利用稀有物种估算物种多样性的指数,其值越高代表群落物种种类越丰富。以上多样性指数的计算参照何芝等[
利用MeA绘制热图,利用Origin2022进行冗余分析(RDA)和绘图,采用SPSS16.0软件进行单因素方差分析(ANOVA),统计学检验方法采用邓肯(Duncan)检验法(
不同处理下土壤脲酶活性整体上随着种植茬数的增加表现出先增后减的变化规律(
不同施肥处理下土壤酶活性随收获茬数的动态变化
Dynamic change characteristics of soil enzyme activity with the harvest number under different fertilization treatments
土壤细菌α多样性指数中仅辛普森指数在各处理中差异显著(
不同施肥处理下土壤细菌α多样性指数
The α diversity index of soil bacteria under different fertilization treatments
处理Treatment | 香农指数Shannon index | 辛普森指数Simpson index | Chao1指数Chao1 index | ACE指数ACE index |
注:ACE:多度覆盖的物种估计量。同列不同小写字母表示不同处理间差异显著( |
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CK | 9.80±0.45a | 0.0054±0.0017ab | 3 349±1 360a | 3 321±274.1a |
KCl | 9.76±0.81a | 0.0050±0.0022ab | 3 249±789.0a | 3 485±721.0a |
KSB | 9.36±0.37a | 0.0101±0.0051a | 2 710±46.09a | 3 725±161.7a |
BC | 10.23±0.46a | 0.0028±0.0009b | 3 200±186.1a | 3 965±176.3a |
BC-KSB | 10.09±0.27a | 0.0036±0.0006ab | 4 061±847.5a | 4 303±1 267a |
解钾菌和生物质炭施用会导致土壤微生物组成发生改变(
不同施肥处理在门水平上对土壤细菌群落组成的影响
Effect of different fertilization treatments on soil bacterial community composition at the level of phylum
根据细菌属水平的丰度结果,采用Z-score标准化方法对丰度排名前30的菌属进行归一化处理并绘制了土壤细菌属水平群落结构组成热图(
生物质炭负载解钾菌对土壤微生物群落结构的影响
Effects of potassium solubilizing bacteria loaded biochar on soil microbial community structure
选择在不同处理土壤样品中相对丰度均超过1 %的土壤细菌优势属进行具体分析,包括8种:罗河杆菌属(
各施肥处理相比CK均显著提升了黑麦草干物质量(
不同施肥处理下第五茬黑麦草干物质量和土壤生物与化学性质
Dry weight of ryegrass and soil biological and chemical properties of the fifth crop under different fertilization treatments
处理Treatment | Y/g | pH | OM /(g·kg–1) | TN/(g·kg–1) | TP /(g·kg–1) | TK/(g·kg–1) | AK /(mg·kg–1) | MBC /(mg·kg–1) | MBN /(mg·kg–1) |
注:Y:黑麦草干物质量;pH:土壤酸碱度;OM:有机质;TN:全氮;TP:全磷;TK:全钾;AK:速效钾;MBC:微生物生物量碳;MBN:微生物生物量氮。下同。Note:Y:Dry biomass of ryegrass;pH:Soil pH;OM:Organic matter;TN:Total nitrogen;TP:Total phosphorus;TK:Total potassium;AK:Available potassium;MBC:microbial biomass carbon;MBN:microbial biomass nitrogen. The same below. | |||||||||
CK | 0.42±0.01e | 4.32±0.16b | 3.54±0.31c | 0.88±0.06c | 0.31±0.01a | 5.79±0.01a | 7.13±1.39c | 53.87±6.21c | 13.72±1.08c |
KCl | 0.68±0.01d | 4.24±0.01b | 3.81±0.51c | 0.89±0.07c | 0.32±0.03a | 6.11±1.34a | 10.37±0.02bc | 71.63±6.47bc | 18.92±2.32b |
KSB | 0.94±0.03c | 4.47±0.22ab | 6.20±0.36b | 0.98±0.04bc | 0.31±0.07a | 6.17±0.51a | 13.71±1.41b | 69.49±3.79bc | 19.92±2.48b |
BC | 1.68±0.07b | 4.59±0.09a | 7.00±1.22b | 1.03±0.05b | 0.34±0.01a | 5.87±0.90a | 26.63±5.08a | 86.96±13.83ab | 27.39±2.11a |
BC-KSB | 2.14±0.16a | 4.46±0.07ab | 11.35±1.89a | 1.18±0.05a | 0.38±0.02a | 6.63±0.45a | 15.72±4.24b | 104.3±11.77a | 30.19±2.51a |
利用冗余分析探讨了最后一茬13种环境因子(脲酶、蔗糖酶、酸性磷酸酶、过氧化氢酶、微生物生物量碳、微生物生物量氮、pH、有机质、全氮、全磷、全钾、速效钾和黑麦草干物质量)与土壤细菌优势属的关系。结果(
土壤细菌群落与土壤生物和化学性质的冗余分析(RDA)
Redundancy analysis(RDA)of soil bacterial community and soil biological and chemical properties
土壤酶活性强弱可反映土壤生化过程强度[
土壤是微生物的生活场所,土壤养分的丰缺状况会影响土壤细菌多样性。本研究发现KSB处理降低了物种的丰富度,却增加了菌群的均匀度(
微生物群落的相对丰度、微生物量的变化以及土壤中的特殊微生物类群(指示生物)可用来间接反映土壤微生态系统的稳定性。本研究添加的微嗜酸寡养单孢菌属于变形菌门(Proteobacteria),变形菌门为本研究土壤中最优势菌门,但KSB和生物质炭的单独施用均降低了变形菌门的相对丰度(
在属分类水平上,罗河杆菌属(
土壤为微生物群落生长繁殖提供微环境,其理化性质又间接影响土壤微生物群落结构的组成[
与空白对照相比,单施生物质炭和解钾菌均可明显提高土壤蔗糖酶、脲酶、酸性磷酸酶和过氧化氢酶活性,而生物质炭负载解钾菌可显著提高该四种酶的活性。同时,生物质炭负载解钾菌也可提高土壤微生物的多样性与均匀度,对土壤细菌群落造成影响,提高土壤有益菌群(绿弯菌门、放线菌门、芽孢杆菌属和慢生根瘤菌属)的丰度,抑制土壤有害菌群的繁殖(变形菌门和罗河杆菌属),显著提高黑麦草产量、微生物生物量碳、微生物生物量氮、有机质、全氮和速效钾含量。冗余分析表明有机质、速效钾、微生物生物量氮、酸性磷酸酶和脲酶是影响土壤细菌群落结构的主要限制因子。可见,生物质炭负载解钾菌对于培肥土壤、改良土壤生态系统具有积极的意义,可以更好地推动生物有机肥的开发利用。
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