热带作物学报

处理Treatment	有机质OM/（g·kg^–1）	铵态氮AN/（mg·kg^–1）	速效磷AP/（mg·kg^–1）	速效钾AK/（mg·kg^–1）	pH	盐度SY/%
CK	12.53±0.31^b	35.90±2.72^d	143.33±4.65^c	246.00±6.19^c	5.40±0.10^b	0.19±0.01^a
LNL41	17.00±1.15^a	59.87±3.44^a	187.67±8.41^a	317.67±6.44^a	6.20±0.10^a	0.12±0.01^c
CM	15.63±0.74^a	49.57±3.76^b	159.67±4.51^b	304.40±9.89^a	5.90±0.30^a	0.15±0.02^b
NS	13.70±0.46^b	42.20±1.95^c	151.07±2.27^bc	267.93±7.14^b	5.43±0.25^b	0.16±0.01^b

处理Treatment	有机质OM/（g·kg^–1）	铵态氮AN/（mg·kg^–1）	速效磷AP/（mg·kg^–1）	速效钾AK/（mg·kg^–1）	pH	盐度SY/%
CK	12.53±0.31^b	35.90±2.72^d	143.33±4.65^c	246.00±6.19^c	5.40±0.10^b	0.19±0.01^a
LNL41	17.00±1.15^a	59.87±3.44^a	187.67±8.41^a	317.67±6.44^a	6.20±0.10^a	0.12±0.01^c
CM	15.63±0.74^a	49.57±3.76^b	159.67±4.51^b	304.40±9.89^a	5.90±0.30^a	0.15±0.02^b
NS	13.70±0.46^b	42.20±1.95^c	151.07±2.27^bc	267.93±7.14^b	5.43±0.25^b	0.16±0.01^b

处理Treatment	叶绿素Chlorophyll/SPAD	穗长Ear length/mm	千粒质量Thousand seed eight/g
CK	30.50±2.69^d	66.79±2.32^c	53.77±1.76^c
LNL41	51.20±1.40^a	79.03±2.15^a	57.23±1.19^a
CM	44.03±2.18^b	73.90±3.22^b	56.50±0.96^ab
NS	38.47±2.30^c	70.93±1.99^bc	54.57±1.01^b

处理Treatment	叶绿素Chlorophyll/SPAD	穗长Ear length/mm	千粒质量Thousand seed eight/g
CK	30.50±2.69^d	66.79±2.32^c	53.77±1.76^c
LNL41	51.20±1.40^a	79.03±2.15^a	57.23±1.19^a
CM	44.03±2.18^b	73.90±3.22^b	56.50±0.96^ab
NS	38.47±2.30^c	70.93±1.99^bc	54.57±1.01^b

处理Treatment	主花期Flower stage	膨大期Expansion stage	灌浆期Grouting stage	成熟期Maturity stage
CK	27.33±4.51^a		45.33±3.06^a		76.00±4.36^a		85.33±2.08^a
LNL41	2.67±0.58^d	90.06±2.86^a	6.33±2.52^d	85.94±5.62^a	11.67±3.21^d	84.50±4.90^a	15.67±3.51^d	81.64±4.02^a
CM	8.33±1.53^c	69.18±6.03^b	20.67±1.53^c	54.19±5.82^b	29.33±3.21^c	61.15±6.66^b	40.33±4.16^c	52.78±3.92^b
NS	14.33±2.52^b	47.19±8.22^c	28.00±3.00^b	38.34±2.62^c	49.67±2.08^b	34.52±4.25^c	66.67±3.51^b	21.91±2.28^c

处理Treatment	主花期Flower stage	膨大期Expansion stage	灌浆期Grouting stage	成熟期Maturity stage
CK	27.33±4.51^a		45.33±3.06^a		76.00±4.36^a		85.33±2.08^a
LNL41	2.67±0.58^d	90.06±2.86^a	6.33±2.52^d	85.94±5.62^a	11.67±3.21^d	84.50±4.90^a	15.67±3.51^d	81.64±4.02^a
CM	8.33±1.53^c	69.18±6.03^b	20.67±1.53^c	54.19±5.82^b	29.33±3.21^c	61.15±6.66^b	40.33±4.16^c	52.78±3.92^b
NS	14.33±2.52^b	47.19±8.22^c	28.00±3.00^b	38.34±2.62^c	49.67±2.08^b	34.52±4.25^c	66.67±3.51^b	21.91±2.28^c

处理Treatment	特征数Feature	ACE	Chao1	Simpson	Shannon	覆盖率Coverage
CK	1706.33±57.95^c	2383.11±51.21^c	2347.26±109.92^b	0.9955±0.00^a	9.1621±0.14^b	0.9498±0.00^a
LNL41	2142.00±3.61^a	2866.46±22.98^a	2788.01±39.11^a	0.9964±0.00^a	9.4859±0.04^a	0.9570±0.00^a
CM	2094.67±74.45^ab	2821.87±49.92^a	2781.99±68.86^a	0.9957±0.00^a	9.2871±0.02^b	0.9606±0.00^a
NS	1963.67±131.56^b	2688.58±32.56^b	2649.82±55.97^a	0.9958±0.00^a	9.2581±0.06^b	0.9567±0.00^a

处理Treatment	特征数Feature	ACE	Chao1	Simpson	Shannon	覆盖率Coverage
CK	1706.33±57.95^c	2383.11±51.21^c	2347.26±109.92^b	0.9955±0.00^a	9.1621±0.14^b	0.9498±0.00^a
LNL41	2142.00±3.61^a	2866.46±22.98^a	2788.01±39.11^a	0.9964±0.00^a	9.4859±0.04^a	0.9570±0.00^a
CM	2094.67±74.45^ab	2821.87±49.92^a	2781.99±68.86^a	0.9957±0.00^a	9.2871±0.02^b	0.9606±0.00^a
NS	1963.67±131.56^b	2688.58±32.56^b	2649.82±55.97^a	0.9958±0.00^a	9.2581±0.06^b	0.9567±0.00^a

绿农林^®41复合微生物菌肥改善连作胡椒土壤细菌群落结构和防控枯萎病的作用

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汪军 , 周游 , 梁昌聪 , 郭立佳 , 杨扬 , 黄俊生 , 杨腊英 ^** , 他永全 ^**

热带作物学报 | 植物保护与生物安全 2025,46(12): 2995-3008

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热带作物学报 |植物保护与生物安全 2025 , 46 (12) : 2995 -3008

绿农林^®41复合微生物菌肥改善连作胡椒土壤细菌群落结构和防控枯萎病的作用

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汪军（1981—），男，博士，助理研究员，研究方向：根际微生态与土传病害

周游（1983—），男，博士，助理研究员，研究方向：植物病害生物防治。

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周游（1983—），男，博士，助理研究员，研究方向：植物病害生物防治。

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汪军, 周游, 梁昌聪, 郭立佳, 杨扬, 黄俊生, 杨腊英^**, 他永全^**

作者信息

中国热带农业科学院环境与植物保护研究所/热带作物生物育种全国重点实验室/农业农村部热带作物有害生物综合治理重点实验室/国家肥料微生物种质资源库（海南）/海南省热带农业微生物菌种资源库，海南海口　571101

通讯作者:

^** 杨腊英（YANG Laying），E-mail：layingyang1980@163.com

他永全（TA Yongquan），E-mail：yongquanta@163.com。

Effects of Lvnonglin ^®41 Compound Microbial Fertilizer Improving Bacterial Community Structure and the Control Effect of Fusarium Wilt Diseases of Continuous Cropping Black Pepper

Jun WANG, You ZHOU, Changcong LIANG, Lijia GUO, Yang YANG, Junsheng HUANG, Laying YANG^**, Yongquan TA^**

Affiliations

Institute of Environment and Plant Protection, Chinese Academy of Tropical Agricultural Sciences / National Key Laboratory for Tropical Crop Breeding / Key Laboratory of Integrated Pest Management on Tropical Crops, Ministry of Agriculture and Rural Affairs / National Collection of Microbial Resource for Fertilizer (Hainan) / Collection of Tropical Agricultural Microbial Resource in Hainan Province, Haikou, Hainan 571101, China

出版时间: 2025-12-25 doi: 10.3969/j.issn.1000-2561.2025.12.017

文章导航

摘要

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从土壤养分和微生物群落结构及多样性角度，研究绿农林^®41复合微生物菌肥改善胡椒连作障碍的机理和效果，为海南胡椒产业化种植提供技术措施。以往年胡椒枯萎病严重发生地块为试验地，分别设置4个处理：清水对照（CK）、绿农林^®41复合微生物菌肥（LNL41）、复合微生物（CM）、菌肥基质（NS），测定根际枯萎病发病率、植株生长和土壤养分；采用16S rDNA测序技术，探究施用LNL41条件下胡椒枯萎病发生及根际土壤细菌群落结构的差异。结果表明：与CK相比，各处理均有一定效果，其中以LNL41处理的效果最为显著。LNL41和CM处理的土壤养分指标均显著高于CK；LNL41、CM和NS处理的叶绿素含量、穗长和千粒质量增幅分别达26.12%~67.87%、6.20%~18.33%和1.48%~6.44%；LNL41处理在不同生长期的发病率达2.67%~15.67%，防效达81.64%~90.06%。根际土壤细菌Ace指数和Chao1指数分别提高了12.82%~20.28%和12.89%~18.78%，Shannon指数提高了1.05%~3.53%，各处理的Simpson指数差异不显著。在目分类水平上，Chitinophagales、Rhizobiales和Burkholderiales是优势细菌目；在属分类水平上，Gaiella、P3OB 42、Lactobacillus、Pseudolabrys和Terrimonas是优势细菌属；各处理共有优势属Bacillus与Candidatus Omnitrophus丰度相似。线性判别分析（LEfSe）结果显示，LNL41处理中存在6个指示菌群；Ellin6067和Tepidisphaera与土壤pH、有机质、速效钾、铵态氮和有效磷呈显著或极显著正相关；网络分析结果进一步表明，LNL41处理能增加土壤细菌共发生网络的复杂性和稳定性；Bugbase功能预测表明，LNL41处理的stress tolerant功能类群丰度增加了5.38个百分点，而CM和NS处理则分别下降了10.43个百分点和7.25个百分点。LNL41能显著改善土壤养分比例，进而改善土壤细菌群落结构和功能特性，激发细菌压力耐受功能，促进胡椒生长，减少枯萎病的发病率。

关键词

绿农林^®41复合微生物菌肥 / 胡椒枯萎病 / 土壤 / 细菌群落 / 结构 / 功能

Abstract

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The purpose of this study was to investigate the mechanism and efficacy of Lvnonglin ^®41 compound microbial fertilizer in mitigating black pepper continuous cropping obstacles from the perspectives of soil nutrients, microbial community structure and diversity, so as to provide technical strategies for the industrial cultivation of black pepper in Hainan. Field experiments were conducted on a plot with a history of severe black pepper Fusarium wilt. Four treatments were designed: water control (CK), Lvnonglin^® 41 compound microbial fertilizer (LNL41), compound microorganisms (CM), and bacterial fertilizer nutrient substrate (NS). The incidence of Fusarium wilt in the rhizosphere, plant growth and soil nutrients were measured. Using 16S rDNA sequencing technology, the differences in the occurrence of black pepper Fusarium wilt and the bacterial community structure in the rhizosphere soil under LNL41 application were explored. The results showed that compared with CK, all treatments exhibited certain effects, with the LNL41 treatment being the most effective. Soil nutrient indicators in the LNL41 and CM treatments were significantly higher than those in CK. The increases in chlorophyll content, spike length and 1000-grain weight under LNL41, CM and NS treatments reached 26.12%–67.87%, 6.20%–18.33% and 1.48%–6.44%, respectively. The incidence rates at different growth stages in the LNL41 treatment were 2.67%–15.67%, with control efficacies of 81.64%–90.06%. The Ace and Chao1 indices of rhizosphere soil bacteria increased by 12.82%–20.28% and 12.89%–18.78%, respectively, and the Shannon diversity index increased by 1.05%–3.53%, while the Simpson index showed no significant difference among treatments. At the order level, Chitinophagales, Rhizobiales and Burkholderiales were the dominant bacterial orders. At the genus level, Gaiella, P3OB 42, Lactobacillus, Pseudolabrys and Terrimonas were the dominant bacterial genera. The abundances of the common dominant genus Bacillus and Candidatus Omnitrophus were similar across treatments. Linear discriminant analysis (LEfSe) results indicated the presence of six indicator bacterial taxa in the LNL41 treatment. Ellin6067 and Tepidisphaera showed significant or highly significant positive correlations with soil pH, organic matter, available potassium, ammonium nitrogen and available phosphorus. Network analysis further revealed that the LNL41 treatment enhanced the complexity and stability of the soil bacterial co-occurrence network. Bugbase functional prediction demonstrated that the abundance of stress tolerant functional groups in the LNL41 treatment increased by 5.38 percentage points, while it decreased by 10.43 and 7.25 percentage points in the CM and NS treatments, respectively. LNL41 significantly improved the ratio of soil nutrients, thereby enhancing the structure and functional characteristics of the soil bacterial community, stimulating bacterial stress tolerance functions, promoting black pepper growth, and reducing the incidence of Fusarium wilt.

Key words

Lvnonglin^® 41 compound microbial fertilizer / black pepper Fusarium wilt / soil / bacteria community / structure / function

引用本文

汪军, 周游, 梁昌聪, 郭立佳, 杨扬, 黄俊生, 杨腊英, 他永全. 绿农林^®41复合微生物菌肥改善连作胡椒土壤细菌群落结构和防控枯萎病的作用. 热带作物学报, 2025 , 46 (12) : 2995 -3008 . DOI: 10.3969/j.issn.1000-2561.2025.12.017

Jun WANG, You ZHOU, Changcong LIANG, Lijia GUO, Yang YANG, Junsheng HUANG, Laying YANG, Yongquan TA. Effects of Lvnonglin ^®41 Compound Microbial Fertilizer Improving Bacterial Community Structure and the Control Effect of Fusarium Wilt Diseases of Continuous Cropping Black Pepper[J]. Chinese Journal of Tropical Crops, 2025 , 46 (12) : 2995 -3008 . DOI: 10.3969/j.issn.1000-2561.2025.12.017

正文

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胡椒（Piper spp.）为多年生木质藤本植物，享有“香料之王”的称号，是全球重要的香辛料作物之一，兼具显著的药用与工业价值^[1]。我国作为世界主要胡椒生产国之一，自1947年引种以来，胡椒已发展成为热区乡村振兴的关键热带经济作物。截至2017年，全国胡椒种植面积接近3万hm²，年产量达3.60万t，产量位居全球第五^[2]。我国胡椒种植具有明显的区域特征，主要集中在海南、云南等地，其中海南是我国胡椒主产区，占全国胡椒种植面积和产量的97%以上，连作现象十分普遍^[3]，连作障碍问题严重，导致产量急剧下降。连作障碍是农业集约化程度提升与复种指数增加共同引发的严重问题，如烟草、黄瓜等作物长期连作会改变土壤理化性质，通过化感物质自毒作用及根际微生物群落的定向演替，引发土壤微生物群落结构失衡。主要表现为细菌型土壤向真菌型土壤转变，特别是镰刀菌数量增多^[4-6]。镰刀菌（Fusarium solani，F. oxysporum Schl. f. sp. piperis）是胡椒枯萎病的主要病原菌^[7-9]，如马来西亚胡椒枯萎病的严重发生导致胡椒植株整株枯萎死亡，胡椒种植园的经济寿命从20 a缩短至6~8 a，造成巨大的经济损失^[10]，枯萎病是胡椒连作障碍中危害严重的真菌性土传病害之一^[3]。为了防止胡椒连作导致的产量下降，椒农常需频繁施用化学农药进行防治，然而，这种做法易引发环境污染、生态失衡和农药残留超标等问题，不利于产区胡椒种植及相关产业的可持续发展。在此背景下，采用微生物制剂（如拮抗菌制剂）可为解决这一问题提供新途径。

目前对胡椒枯萎病的研究主要集中在病原菌的分离鉴定、抗病性、活性成分和盆栽防效等方面^[10-13]，而关于微生物制剂改善胡椒根际土壤细菌群落结构变化和防控胡椒枯萎病的相关报道较少，但在其他作物上已有大量研究，多功能微生物菌群与特定营养基质复配制成的微生物菌肥，在防控土传病害、提升作物产量和品质并缓解连作障碍方面具有广阔前景^[14-16]。含有木霉菌（Trichoderma spp.）和芽孢杆菌（Bacillus spp.）等功能菌株和有机营养制备的生物有机肥，具有增加土壤肥力、增强作物抗病能力、改善连作土壤理化性状、优化土壤微生物菌群的作用，富集了Bacillus、Ammoniphilus、Acidibacter等具有溶矿功能的细菌，降低了拮抗微生物的活性并降低了镰刀菌（Fusarium）、莱氏菌（Lectera）等病原菌丰度，同时增加了具有抗真菌活性的土著微生物如溶血杆菌属（Lysobacter spp.）的丰度^[17-23]。生物菌肥驱动土壤中有益微生物成为微生物网络的关键类群^[24]，例如WANG等^[25]通过轮作菠萝结合生物有机肥防控香蕉枯萎病的研究发现，土壤中的细菌结构、真菌结构和伯克霍尔德菌（Burkholderia）相对丰度对病害发生率的约束作用最为显著。因此，研究微生物菌肥对根际微生物群落和连作枯萎病发生的影响，对保障胡椒产业健康发展至关重要。。

绿农林^®41复合微生物菌肥（LNL41）为本团队研发并正式登记的产品，由优良生防菌长枝木霉（T. longibrachiatum H02）、专利菌株绿色木霉（T. viride H06）、枯草芽孢杆菌（B. subtilis BLG010）与氨基酸等营养载体复配制成。本研究团队前期已对LNL41在香蕉和番茄上的应用进行了报道^[26-27]，而有关LNL41对胡椒连作土壤理化性质、根际微生物群落的影响和枯萎病的防治效果缺乏深入研究。为此，本研究在连作胡椒枯萎病高发地块展开试验，设置LNL41处理组，研究其对土壤养分、植株生长及病害发生率的影响，并利用16S rDNA测序技术分析胡椒根际细菌群落结构的变化。旨在探明LNL41与胡椒枯萎病发生、根际土壤理化特性及细菌群落间的关联机制，为有效防控连作胡椒枯萎病提供理论支撑。

1　材料与方法

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1.1　材料

供试菌株：生防菌长枝木霉H02、专利菌株绿色木霉H06和枯草芽孢杆菌BLG010由中国热带农业科学院环境与植物保护研究所提供。

供试复合物微生物（CM）：由生防菌长枝木霉H02、专利菌株绿色木霉H06（ZL2012103 23738.2）和枯草芽孢杆菌BLG010（ZL2012102 48300.2）分别经活化、发酵、离心、去除代谢物，收集沉淀的高浓度菌体并干燥复配获得干粉，总有效活菌数为5×10⁹ CFU/g，其中H02活菌数为8×10⁸ CFU/g，H06活菌数为14×10⁸ CFU/g，BLG010活菌数为28×10⁸ CFU/g。

供试氨基酸粉：氨基酸含量为35.6%，由四川世宏科技有限公司生产。

供试绿农林^®41复合微生物菌肥（LNL41），微生物肥（2019）准字（7527）号，由中国热带农业科学院环境与植物保护研究所研制，海南宝绿春农业开发有限公司生产。取长枝木霉H02、专利菌株绿色木霉H06和枯草芽孢杆菌BLG010的高浓度菌体干粉、氨基酸粉和复合肥（15-15-15）复配制备获得。有效成分包括：有机质含量≥40%，总养分（N+P₂O₅+K₂O）为12%，总有效活菌数为5×10⁹ CFU/g，其中H02活菌数为8×10⁸ CFU/g，H06活菌数为14×10⁸ CFU/g，BLG010活菌数为28×10⁸ CFU/g。

供试菌肥营养基质（NS）：取LNL41水溶后，在121 ℃，20 min条件下湿热灭菌。养分指标与LNL41相同[有效成分包括：有机质含量≥40%，总养分（N+P₂O₅+K₂O）为12%]。

供试胡椒品种：大叶胡椒。

1.2　方法

1.2.1　试验地点

试验时间为2024年9月10日至2025年7月10日；试验地点设在海南省文昌市文城镇新合村胡椒基地，连作年限达13 a，土壤较为板结，酸化现象较为严重，肥力有所下降，枯萎病发生率呈上升趋势，产量较低。常年胡椒枯萎病发生严重。土壤以砖红壤为主，土体深厚，质地多为砂质壤土。试验开始前耕层土壤基本理化性质：有机质含量为11.07 g/kg，铵态氮含量为34.48 mg/kg，速效磷含量为135.67 mg/kg，速效钾含量为231.32 mg/kg，pH为5.48。

1.2.2　试验设计

田间试验在常规施肥的基础上，分别设置4个处理：清水对照（CK）、绿农林^®41复合微生物菌肥（LNL41）、菌肥营养基质（NS）和复合微生物（CM）。不同处理随机区组排列，每处理各100株，每处理3个重复。自2024年9月10日开始，施用时分别取LNL41、CM和NS各5 kg用2500 kg清水稀释至500倍，浇灌于胡椒根部，每间隔3周浇灌1次，共13次。

1.2.3　土壤养分测定

在胡椒采收期，每个重复采用五点取样法选取5株胡椒的10~20 cm根围土壤200 g，混匀、去除石块和植物残体、风干，过200目筛后，运用灼烧法测量土壤有机质含量，利用HM-TYA型土壤肥料养分速测仪测定土壤速效氮、速效磷、速效钾含量，利用TR-8D盐度检测仪测定土壤盐度，利用pH计测定土壤酸碱度。

1.2.4　胡椒生长测定和田间病害调查

在胡椒成熟期，自然光下分别从每株上、下部选取健康叶片，用叶绿素测定仪TYS-4N测定叶片叶绿素含量（SPAD值）；参照祖超等^[28]的方法测定千粒质量、穗长。参照文献[8-9]的方法调查胡椒枯萎病在主花期（2024年10月20日）、膨大期（2025年1月20日）、灌浆期（2025年3月20日）和成熟期（2025年6月20日）的发病率，计算不同处理对胡椒枯萎病的防治效果。发病率=发病株数/调查总株数×100%；防治效果=（对照发病率–处理发病率）/对照发病率×100%。

1.2.5　土壤细菌微生物多样性测定

取适量1.2.3中制备的土样装入灭菌离心管中，置于–80 ℃超低温冰箱中速冻后，委托北京百迈客生物科技有限公司检测土壤细菌多样性。采用TGuide S96磁珠法土壤/粪便基因组DNA提取试剂盒提取土壤总DNA，基于PacBio测序平台，利用单分子实时测序（single molecule real-time sequencing）的方法对marker基因进行测序。

1.3　数据处理

采用Excel 2016、SAS 9.0软件对试验数据进行统计分析，采用Duncan氏新复极差法进行差异显著性检验。使用QIIME2软件、R语言对测序结果进行样品α、β多样性、物种丰度和聚类热图分析，基于效应大小线性判别（linear discriminant analysis effect size，LEfSe）方法分析差异物种，用Pearson法对微生物、土壤养分指标进行相关性分析，采用BugBase软件分析细菌表型，以上分析在百迈克云平台（https://international.biocloud.net）上完成。根据ggClusterNet函数分析土壤细菌分子生态网络。

2　结果与分析

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2.1　不同处理对胡椒根际土壤养分含量的影响

与CK相比，各处理对土壤养分的改善作用依次表现为LNL41>CM>NS。各处理间铵态氮含量差异显著；CK和CM处理的各指标均差异显著；CK和NS处理的有机质、速效磷含量和pH差异不显著，其余指标间差异显著；LNL41和CM处理的有机质、速效钾含量和pH差异不显著，其余指标差异显著；LNL41和NS处理的各指标均差异显著；CM和NS处理的速效磷含量和盐度差异不显著，其余指标差异显著（表1）。

2.2　不同处理对胡椒生长和发病率的影响

通过调查不同处理对成熟期胡椒生长的影响发现，与CK相比，各处理的促生作用表现为LNL41>CM>NS，各处理的叶绿素含量、穗长和千粒质量增幅分别达26.12%~67.87%、6.20%~18.33%和1.48%~6.44%。各处理的叶绿素含量差异显著；CK、LNL41和CM处理的穗长差异显著，而CK与NS、CM与NS之间的穗长差异不显著；CK、LNL41和NS处理的千粒质量差异显著，而LNL41与NS、CM与NS处理的千粒质量差异不显著（表2）。

通过调查不同处理对不同生育期胡椒枯萎病发病率的影响发现，与CK相比，各处理的控病作用表现为LNL41>CM>NS。在主花期、膨大期、灌浆期和成熟期各处理的发病率和防效均达到显著差异，4个时期的发病率分别为2.67%~14.33%、6.33%~28.00%、11.67%~49.67%和15.67%~66.67%，防效分别达47.19%~90.06%、38.34%~85.94%、34.52%~84.50%和21.91%~81.64%（表3）。

2.3　不同处理连作胡椒根际细菌α、β多样性分析

根据表4可知，各处理的覆盖率（coverage）无显著差异，范围为0.9498~0.9606，说明各处理的样本OTU覆盖度已经饱和，未检到基因序列的概率很低，本次测序结果能够代表不同处理方式下连作胡椒根际土壤细菌群落的真实情况。

与CK相比，LNL41、CM和NS处理的物种丰度指标特征数、ACE指数、Chao1指数和Shannon指数分别提高了15.08%~25.53%、12.82%~20.28%、12.89%~18.78%和1.05%~3.53%；CK与LNL41处理的特征数、ACE指数、Chao1指数和Shannon指数差异显著；CK与CM、CK与NS处理的特征数Feature、ACE指数和Chao1指数差异显著；LNL41与CM处理的特征数、ACE指数和Chao1指数差异不显著；LNL41与NS处理的特征数、ACE指数和Shannon指数差异显著；CM与NS处理仅ACE指数差异显著；各处理的Simpson指数差异不显著（表4）。

在OTU水平对不同处理细菌群落进行PCoA分析，其中PC1的贡献率为31.52%，PC2的贡献率为14.15%，二者累积贡献率为45.67%（图1）。3个处理能明显分开，说明相同处理样品的细菌群落结构差异较小，不同处理样品的细菌群落结构差异较大。

2.4　目和属水平上细菌群结构组成分析

目水平丰度相对含量（OTUs数相对丰度前15）排名前5的细菌目中，Solirubrobacterales为CK的独有优势目；Rhizobiales和Burkholderiales为各处理的共有优势目；Lactobacillales为CK、LNL41和CM处理的共有优势目；Gaiellales为CK、LNL41和NS处理的共有优势目；Chitinophagales为LNL41、CM和NS处理的共有优势目；Myxococcales为CM和NS处理的共有优势目；Bacillales在CK、LNL41、CM和NS各处理中的丰度排名分别为14（1.61%）、12（2.19%）、13（1.39%）、15（1.58%）（图2A）。

进一步对各分组样品在属水平（OTUs数相对丰度前15）排名前5的相对丰度进行分析，结果表明，不同处理的细菌群落结构存在明显差异，排名前5的细菌属中，unclassified Micropepsaceae为CK独有优势属；Gaiella为CK、LNL41和NS处理的共有优势属；P3OB_42为各处理的共有优势属；Lactobacillus为CK和LNL41处理的共有优势属；Terrimonas为LNL41、CM和NS处理的共有优势属；Nitrospira为LNL41和CM处理的共有优势属；Pseudolabrys为CK和CM处理的共有优势属；Pedomicrobium分别为CK和NS的独有优势属；Bacillus在CK、LNL41、CM和NS各处理中的丰度排名分别为9（1.50%）、9（2.12%）、15（1.24%）、12（1.49%）（图2B）。

2.5　物种丰度聚类热图

在物种丰度聚类热图中，横向聚类表明，不同物种在CK、LNL41和CM处理的枝长较短，3个处理间丰度相似；纵向聚类显示，各处理共有优势属Bacillus与Candidatus Omnitrophus丰度相似；Haliangium和Pedomicrobium分别与Lactobacillus和Pseudolabrys相似（图3）。

2.6　差异物种分析

基于效应大小线性判别分析（linear discriminant analysis effect size，LEfSe）方法，分析不同处理属水平的显著性差异的标志性物种（图4A）。通过设置线性判别分析（linear discriminant analysis，LDA，LDA≥3.5），4个处理共筛选出41个biomarkers（图4B）。与CK比较，LNL41处理有Limosilactobacillus、Ellin6067、Ramlibacter、unclassified Chitinophagales、unclassified Prolixibacteraceae、unclassified Steroidobacteraceae共6类物种显著高于其他处理；CM处理有unclassified Entotheonellaceae、Nitrospira、RB41、P3OB_42、uncultured Candidatus Rokubactera bacterium、Flavobacterium、AKYG587、Tepidisphaera共8类物种显著高于其他处理；NS处理有Terimonas、Pedomicrobium、Candidatus Omnitrophus、Gaiella、unclassified TRA3_20、Subgroup_10、unclassified Ilumatobacteraceae、Povalibacter共8类物种显著高于其他处理。

2.7　根际土壤微生物与环境因子的相关性分析

根际土壤细菌（丰度排名前15）中，Ellin6067和Tepidisphaera与土壤pH、有机质、速效钾、铵态氮和有效磷呈显著或极显著正相关，Gaiella和unclassified Gaiellales与上述土壤指标呈显著或极显著负相关；Tepidisphaera与土壤盐度呈极显著负相关，而unclassified Gaiellales与土壤盐度呈极显著正相关（图5）。

2.8　不同处理对土壤细菌分子生态网络的影响

基于属水平进行不同处理微生物共发生网络分析。结果表明，不同处理的微生物共发生网络中微生物组成较为相似，均有酸杆菌门（Acidobacteriota）、放线菌门（Actinobacteriota）、拟杆菌门（Bacteriodota）和蛭弧菌门（Bdellovibrionota）等13个门水平下的微生物组成（图6）。然而，不同处理的微生物共发生网络中微生物成员之间的互作关系具有较大的差异。与其他处理相比，LNL41处理的网络节点数、边数、正相关边数以及网络平均度等参数均有所增加，表明LNL41处理能增加土壤细菌共发生网络的复杂性和稳定性。并且该结果与LNL41处理的微生物多样性增大的结果一致。

2.9　土壤细菌群落功能预测

基于BugBase数据库对胡椒根际土壤细菌表型进行预测分析，共检测到9种微生物表型（图7）。与CK相比，总体表现为LNL41处理的stres tolerant功能类群丰度增加了5.38个百分点，而CM和NS处理的则分别下降了10.43个百分点和7.25个百分点。具体表现为：LNL41处理的gram negative、potentially pathogenic、stress tolerant和anaerobic的功能类群丰度增幅较大，达到4.31%~9.83%；CM处理的gram negative、potentially pathogenic、anaerobic功能类群丰度增幅较大，达到7.66%~11.78%；NS处理的potentially pathogenic和anaerobic功能类群丰度增幅较大，达到4.13%~4.96%。与CK相比，LNL41处理的forms biofilms、gram positive、contains mobile elements、aerobic功能类群丰度下降幅度为5.55%~11.92%；CM处理的stress tolerant、contains mobile elements、gram positive、aerobic功能类群丰度降幅较大，达到10.43%~17.44%；NS处理的facultatively anaerobic、stress tolerant、aerobic功能类群丰度降幅较大，达到3.31%~4.30%。

3　讨论

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微生物菌肥兼具改善土壤理化性质（如养分状况）、调节根际微生态平衡、抑制土传病害以及促进作物增产提质等多重功能。本项目组研究发现氨基酸作为营养载体促进芽孢杆菌菌体增殖、分泌抑菌物质和促生长物质^{[27, 29]}。ZHENG等^[30]研究发现土壤养分和细菌群落失衡是土传枯萎病的重要特征。YU等^[31]研究发现微生物肥料能有效缓解土壤酸化，增加土壤速效钾和有机质含量，并有利于提高根际土壤硝酸还原酶活性，在提高土壤微生态方面具有重要作用。本研究应用含有长枝木霉、绿色木霉和枯草芽孢杆菌作为功能菌株的复合微生物菌肥LNL41，明显提升了胡椒根际土壤的有机质、铵态氮、速效磷、速效钾含量和pH，降低了盐度，显著优于CM和NS处理，与YU等^[31]和李得铭等^[32]应用微生物肥缓解土壤酸化、提高土壤速效钾和有机质含量的研究结果相似，由此推测微生物菌肥中的功能菌株芽孢杆菌和木霉通过代谢氨基酸类营养物质实现高效增殖，进而活化根际难溶性矿物养分，显著提升土壤养分含量。

汪军等^[26]研究表明，施用LNL41微生物菌肥增加了番茄株高、茎围、叶面积、光合作用、叶绿素、产量和水溶性糖；ZHOU等^[14]施用含有木霉和芽孢杆菌的微生物制剂显著促进了番茄SPAD值、防御酶活性、D-果糖等指标的提升。本研究也发现促生作用表现为LNL41>CM>NS，千粒质量、叶绿素含量、穗长均增幅明显，表现出对连作胡椒的增产作用，其原因可能是LNL41提高了胡椒根系的养分吸收效率，以及对有害生物和非生物因子的耐受力，从而促进其生长。微生物菌肥制剂具有改善土壤生态环境、抑制病原菌增殖、提高作物免疫力、增加有益微生物数量等功效，能有效抑制土传病害的发生^{[26, 32-34]}。CHEN等^[35]发现施用含有地衣芽孢杆菌（Bacillus licheniformis X-1）和甲基营养型芽孢杆菌（B. methylotrophicus Z-1）的生物有机肥后，草莓枯萎病发病率降低82.84%；汪军等^{[16, 26]}应用含有木霉、淡紫拟青霉和枯草芽孢杆菌的微生物菌肥后，番茄和香蕉枯萎病发病率分别下降至16.33%和1.10%。本研究中，LNL41对胡椒各生长期的枯萎病发病率控制在2.67%~15.67%之间，防效维持在81.64%~90.06%之间，而单独施用CM和NS处理的从苗期至成熟期其防效均大幅下降，成熟期防效仅分别为21.91%和52.78%，表明复合微生物菌群需依赖营养基质中的关键物质完成增殖过程并分泌拮抗活性成分，有效重构了根际微生态环境，增强了根系抗病能力。

施用木霉、芽孢杆菌等生防制剂可增加根际土壤等有益微生物的种群丰度，诱导土壤细菌群落丰富度和多样性增加来抑制枯萎病等土传病害^{[17, 35-36]}。本研究中施用LNL41、NS和CM后的物种丰度指标特征数提高了15.08%~25.53%，丰富度指标ACE指数、Chao1指数分别提高了12.82%~20.28%和12.89%~18.78%，多样性指标Shannon指数提高了1.05%~3.53%；由此推测上述3种处理在抑制病原菌的同时，也提高了土壤中细菌种类的丰度。TANG等^[37]施用含有红杆菌（Erythrobacter sp. YH-07）的有机粪肥BF防治番茄枯萎病，增加了番茄根际微生物丰富度和多样性，以及芽孢杆菌属（Bacillus）、交替红色杆菌属（Altererythrobacte）等有益菌丰度。JAMIL等^[38]发现健康香蕉和感染枯萎病香蕉根际细菌群落关键类群和土壤理化性质的变化对香蕉健康指标具有潜在的指示作用。本研究发现LNL41处理后Bacillales和Bacillus丰度分别为2.19%和2.12%，高于其他处理，Bacillus与Candidatus Omnitrophus丰度相似，同时Limosilactobacillus等6类差异物种显著高于其他处理；Ellin6067和Tepidisphaera与土壤pH、有机质、速效钾、铵态氮和有效磷含量呈正相关；LNL41处理提高了土壤细菌共发生网络的复杂性和稳定性，说明施用LNL41后通过增加胡椒根际土壤细菌的丰度、丰富度和多样性而成为土壤微生物群落组成的主要驱动力，可能有助于诱导胡椒根际有效抵抗病原菌。

BugBase是一种预测复杂微生物组内功能途径的生物水平覆盖以及生物可解释表型的方法。BugBase通过表型情况预测微生物群落生态功能^[39]。WANG等^[40]研究发现，含有芽孢杆菌的微生物有机肥能显著改变烟草根际土壤中细菌功能类群gram-positive、potentially pathogenic、aerobic和gram-negative。YANG等^[41]研究发现，施用多粘类芽孢杆菌（Paenibacillus polymyxa）KN-03菌株能抑制柑橘黄龙病病菌增殖，同时提高biofilm formation、element mobilization和stress tolerant类群的相对丰度。LI等^[42]发现对炭疽病和根腐病具有耐病性的攸县油茶（Camellia yuhsienensis）根际细菌优势功能类群为gram-negative。本研究通过BugBase功能预测得知，与CK比较，LNL41处理的gram negative、potentially pathogenic、stress tolerant和anaerobic功能类群显著增加，而CM和NS处理的stress tolerant功能类群均显著下降；LNL41处理的stress tolerant功能类群丰度增加了5.38个百分点，而CM和NS处理则分别下降了10.43个百分点和7.25个百分点。推测LNL41处理可能通过提高胡椒根际耐逆性细菌丰度，从而有效抑制病原菌增殖。需要注意的是各处理的potentially pathogenic功能类群数量增加，此类细菌表型变化对真菌病害的影响尚不清楚，需进一步研究。今后应进一步探索其生防机理，发掘具有较强抗枯萎病作用的有益微生物，使其在胡椒枯萎病绿色防控领域发挥更好的作用。

4　结论

收起

绿农林^®41复合微生物菌肥可降低连作胡椒枯萎病发病率，提高千粒质量、穗长和叶绿素含量等指标，改善胡椒根际养分有机质、铵态氮、速效磷、盐度和pH，改善土壤细菌群落结构，提升压力耐受功能类群等丰度，表明根际施用绿农林^®41复合微生物菌肥防病促生作用显著，应用前景良好。

基金

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海南省重点研发项目(ZDYF2024YJGG001-25)
海南省胡椒产业技术体系病虫草害防控岗位专家项目(HNARS-09-G03)
中央级公益性科研院所基本科研业务费专项(1630042024002)

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文献

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2025年第46卷第12期

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文章信息

doi: 10.3969/j.issn.1000-2561.2025.12.017

接收时间：2025-08-25
首发时间：2026-06-24
出版时间：2025-12-25

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出版历史

收稿日期：2025-08-25
录用日期：2025-10-09

基金

海南省重点研发项目(ZDYF2024YJGG001-25)

海南省胡椒产业技术体系病虫草害防控岗位专家项目(HNARS-09-G03)

中央级公益性科研院所基本科研业务费专项(1630042024002)

作者信息

通讯作者:

^** 杨腊英（YANG Laying），E-mail：layingyang1980@163.com

他永全（TA Yongquan），E-mail：yongquanta@163.com。

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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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处理Treatment	有机质OM/（g·kg^–1）	铵态氮AN/（mg·kg^–1）	速效磷AP/（mg·kg^–1）	速效钾AK/（mg·kg^–1）	pH	盐度SY/%
CK	12.53±0.31^b	35.90±2.72^d	143.33±4.65^c	246.00±6.19^c	5.40±0.10^b	0.19±0.01^a
LNL41	17.00±1.15^a	59.87±3.44^a	187.67±8.41^a	317.67±6.44^a	6.20±0.10^a	0.12±0.01^c
CM	15.63±0.74^a	49.57±3.76^b	159.67±4.51^b	304.40±9.89^a	5.90±0.30^a	0.15±0.02^b
NS	13.70±0.46^b	42.20±1.95^c	151.07±2.27^bc	267.93±7.14^b	5.43±0.25^b	0.16±0.01^b

处理Treatment

有机质OM/（g·kg^–1）

铵态氮AN/（mg·kg^–1）

速效磷AP/（mg·kg^–1）

速效钾AK/（mg·kg^–1）

盐度SY/%

12.53±0.31^b

35.90±2.72^d

143.33±4.65^c

246.00±6.19^c

5.40±0.10^b

0.19±0.01^a

LNL41

17.00±1.15^a

59.87±3.44^a

187.67±8.41^a

317.67±6.44^a

6.20±0.10^a

0.12±0.01^c

15.63±0.74^a

49.57±3.76^b

159.67±4.51^b

304.40±9.89^a

5.90±0.30^a

0.15±0.02^b

13.70±0.46^b

42.20±1.95^c

151.07±2.27^bc

267.93±7.14^b

5.43±0.25^b

0.16±0.01^b

处理Treatment	叶绿素Chlorophyll/SPAD	穗长Ear length/mm	千粒质量Thousand seed eight/g
CK	30.50±2.69^d	66.79±2.32^c	53.77±1.76^c
LNL41	51.20±1.40^a	79.03±2.15^a	57.23±1.19^a
CM	44.03±2.18^b	73.90±3.22^b	56.50±0.96^ab
NS	38.47±2.30^c	70.93±1.99^bc	54.57±1.01^b

处理Treatment

叶绿素Chlorophyll/SPAD

穗长Ear length/mm

千粒质量Thousand seed eight/g

30.50±2.69^d

66.79±2.32^c

53.77±1.76^c

LNL41

51.20±1.40^a

79.03±2.15^a

57.23±1.19^a

44.03±2.18^b

73.90±3.22^b

56.50±0.96^ab

38.47±2.30^c

70.93±1.99^bc

54.57±1.01^b

处理Treatment	主花期Flower stage	膨大期Expansion stage	灌浆期Grouting stage	成熟期Maturity stage
CK	27.33±4.51^a		45.33±3.06^a		76.00±4.36^a		85.33±2.08^a
LNL41	2.67±0.58^d	90.06±2.86^a	6.33±2.52^d	85.94±5.62^a	11.67±3.21^d	84.50±4.90^a	15.67±3.51^d	81.64±4.02^a
CM	8.33±1.53^c	69.18±6.03^b	20.67±1.53^c	54.19±5.82^b	29.33±3.21^c	61.15±6.66^b	40.33±4.16^c	52.78±3.92^b
NS	14.33±2.52^b	47.19±8.22^c	28.00±3.00^b	38.34±2.62^c	49.67±2.08^b	34.52±4.25^c	66.67±3.51^b	21.91±2.28^c

处理Treatment

主花期Flower stage

膨大期Expansion stage

灌浆期Grouting stage

成熟期Maturity stage

发病率Disease incidence/%

防效Control efficiency/%

发病率Disease incidence/%

防效Control efficiency/%

发病率Disease incidence/%

防效Control efficiency/%

发病率Disease incidence/%

防效Control efficiency/%

27.33±4.51^a

45.33±3.06^a

76.00±4.36^a

85.33±2.08^a

LNL41

2.67±0.58^d

90.06±2.86^a

6.33±2.52^d

85.94±5.62^a

11.67±3.21^d

84.50±4.90^a

15.67±3.51^d

81.64±4.02^a

8.33±1.53^c

69.18±6.03^b

20.67±1.53^c

54.19±5.82^b

29.33±3.21^c

61.15±6.66^b

40.33±4.16^c

52.78±3.92^b

14.33±2.52^b

47.19±8.22^c

28.00±3.00^b

38.34±2.62^c

49.67±2.08^b

34.52±4.25^c

66.67±3.51^b

21.91±2.28^c

处理Treatment	特征数Feature	ACE	Chao1	Simpson	Shannon	覆盖率Coverage
CK	1706.33±57.95^c	2383.11±51.21^c	2347.26±109.92^b	0.9955±0.00^a	9.1621±0.14^b	0.9498±0.00^a
LNL41	2142.00±3.61^a	2866.46±22.98^a	2788.01±39.11^a	0.9964±0.00^a	9.4859±0.04^a	0.9570±0.00^a
CM	2094.67±74.45^ab	2821.87±49.92^a	2781.99±68.86^a	0.9957±0.00^a	9.2871±0.02^b	0.9606±0.00^a
NS	1963.67±131.56^b	2688.58±32.56^b	2649.82±55.97^a	0.9958±0.00^a	9.2581±0.06^b	0.9567±0.00^a

处理Treatment

特征数Feature

ACE

Chao1

Simpson

Shannon

覆盖率Coverage

1706.33±57.95^c

2383.11±51.21^c

2347.26±109.92^b

0.9955±0.00^a

9.1621±0.14^b

0.9498±0.00^a

LNL41

2142.00±3.61^a

2866.46±22.98^a

2788.01±39.11^a

0.9964±0.00^a

9.4859±0.04^a

0.9570±0.00^a

2094.67±74.45^ab

2821.87±49.92^a

2781.99±68.86^a

0.9957±0.00^a

9.2871±0.02^b

0.9606±0.00^a

1963.67±131.56^b

2688.58±32.56^b

2649.82±55.97^a

0.9958±0.00^a

9.2581±0.06^b

0.9567±0.00^a