梨火疫病菌与梨腐烂病菌对生长和致病的相互影响

doi:10.16420/j.issn.0513-353x.2024-0134

园艺学报 ›› 2024, Vol. 51 ›› Issue (11): 2668-2684.doi: 10.16420/j.issn.0513-353x.2024-0134

梨火疫病菌与梨腐烂病菌对生长和致病的相互影响

陈春桥¹, 孙丽英², 盛强³, 韩剑¹, 傅本重¹, 罗明¹^,^*()

¹ 新疆农业大学农学院，农林有害生物检测与防控重点实验室，农业农村部西北荒漠绿洲农林外来入侵生物防控重点实验室，乌鲁木齐 830052
² 西北农林科技大学植物保护学院，陕西杨凌 712100
³ 新疆巴音郭楞蒙古自治州农业科学研究院，新疆库尔勒 841003

收稿日期:2024-04-04 修回日期:2024-07-16 出版日期:2024-12-12 发布日期:2024-11-26
通讯作者:
^* E-mail：luomingxjau@sina.com
基金资助:
新疆维吾尔自治区自然科学基金重点项目(2021D01D12); 国家重点研发计划项目(2021YFD1400200)

Influences of Interactions Between Erwinia amylovora and Valsa pyri on Growth and Pathogenicity

CHEN Chunqiao¹, SUN Liying², SHENG Qiang³, HAN Jian¹, FU Benzhong¹, LUO Ming¹^,^*()

¹ Key Laboratory of Detection and Prevention and Control of Agricultural and Forestry Pests，Key Laboratory of Prevention and Control of Invasive Organisms in Agroforestry in Northwest Desert Oasis，Ministry of Agriculture and Ministry of Agriculture and Rural Affairs，College of Agriculture，Xinjiang Agricultural University，Urumqi 830052，China
² School of Plant Protection，Northwest Agroforestry and Science and Technology University，Yangling，Shaanxi 712100，China
³ Research Institute of Agricultural Science of Bayingolin Mongolian Autonomous Prefecture，Korla，Xinjiang 841003，China

Received:2024-04-04 Revised:2024-07-16 Published:2024-12-12 Online:2024-11-26

摘要/Abstract

摘要：

梨火疫病和梨腐烂病是当前严重危害库尔勒香梨生产的两大病害，在田间常混合发生，加重了危害程度和防治难度。为了明确梨火疫病病原细菌Erwinia amylovora（E.a）和梨腐烂病病原真菌Valsa pyri（V.p）之间的相互作用，探讨复合侵染机制，将梨火疫病菌E. a001菌株（梨腐烂病菌V.pKRL菌株）与V.pKRL（E.a001）的发酵滤液共培养，研究其互作对其生长和致病力的影响。结果表明，与发酵滤液共培养能显著提高两种菌体的生长速率，E.a001活菌数显著高于对照，生物膜形成量是对照的2.2倍，V.pKRL菌丝生物量是对照的1.46倍，分生孢子萌发率提高14.9%。两种发酵滤液共培养均能降低培养液的pH值，而E.a001与梨黑斑病菌（Alternaria alternata）发酵滤液，V.pKRL与梨花枯病菌（Pseudomonas syringae）和梨锈水病菌（Dyckeya fangzhongda sp. nov）发酵滤液共培养则无促进生长和降低pH等作用。E.a001与V.pKRL发酵滤共培养显著增加了E.a001的胞外多糖和梨火疫毒素产量，hrp基因簇中HrpA、hrpB、hrpD、hrpJ、HrpW、hrpY、hrcJ的相对表达量上调1.53 ~ 2.27倍，其中hrpY的上调幅度最大；同样地，V.pKRL与E.a001发酵滤液共培养增加了V. pyri根皮苷降解毒素的合成量，细胞壁降解酶相关基因PG、β-Gal、PME和α-Af相对表达量上调1.64 ~ 1.91倍，其中PG的上调幅度最大。通过对E.a001和V.pKRL单菌接种、同位点和异位点复合接种，比较病原菌的致病力。结果显示，在库尔勒香梨枝条、离体叶片和果实上均表现出同位点复合接种处理的致病力最强，显症时间短，病斑扩展速度、病斑面积、V.pKRL的产孢数均大于单菌接种、异位点复合接种。本研究结果表明，梨火疫病菌和梨腐烂病菌能相互促进生长，增加毒素产生水平和上调主要致病基因表达量，复合侵染增强了致病力，二者存在着特定的协同互作关系。

关键词: 梨, 梨火疫病菌, 梨腐烂病菌, 复合侵染, 细菌—真菌互作, 致病力

Abstract:

Fire blight and pear Valsa canker，two prominent diseases，currently inflict substantial damage on the production of Korla fragrant pear（Pyrus sinkiangensis）. Their often co-occurrence in the field，exacerbating the severity of the diseases and complicating control efforts. In order to clarify the interaction between the fire blight pathogen bacterium（Erwinia amylovora，E.a）and the pear Valsa canker pathogen fungus（Valsa pyri，V.p），and to explore the complex infection mechanisms of these pathogens，the present study was conducted to investigate the effects of pathogen interactions on the growth and pathogenicity of E.a and V.p by using E.a strain E.a001 and V.p strain V.pKRL fermentation filtrates co-culture. The results showed that co-culture of E.a001 with V.pKRL fermentation filtrates significantly enhanced the growth rate of E.a001，with a higher viable cell count compared to the control，and a 2.2-fold increase in biofilm formation compared to the control. Co-culture of V.pKRL with E.a001 frmentation filtrates resulted in a 1.46-fold increase in V.pKRL mycelial biomass，and a 14.9% increase in conidial germination rate compared to the control. Additionally，both co-culture of E.a001 with V.pKRL and V.pKRL with E.a001 fermentation filtrates reduced the pH of the culture medium，whereas co-culture of E.a001 with the reference strains of Alternaria alternata，and V.pKRL with Pseudomonas syringae and Dyckeya fangzhongda sp. nov did not exhibit growth promotion or pH reduction characteristics. It was found that co-culture of E.a001 + V.pKRL fermentation filtrate increased the extracellular polysaccharide and pear fire blight toxin production of E.a001. The relative expression of HrpA，hrpB，hrpD，hrpJ，HrpW，hrpY and hrcJ in the hrp gene cluster was up-regulated by 1.53 to 2.27-fold，with the greatest up-regulation of the hrpY gene. Similarly，E.a001 fermentation filtrate increased the synthesis of V.pKRL phlorizin degradation toxins，and the relative expressions of cell wall degradation-related enzyme genes，PG，β-Gal，PME and α-Af were up-regulated by 1.64 to 1.91-fold，with the largest up-regulation of the PG gene. Pathogenicity assessments on Korla fragrant pear ranches，leaves，and fruits revealed that sequential composite inoculation of E.a001 and V.pKRL at the same site induces the most potent pathogenicity，marked by rapid symptom development，lesion expansion，and increased V.p spore production compared to single or sequential inoculations at different sites. These findings underscore mutual promotion between E.a and V.p，enhancing toxin production，up-regulating pathogenic gene expression，and reinforcing pathogenicity，thereby indicating a specific cooperative interactions relationship between the two pathogens.

Key words: pear, Erwinia amylovora, Valsa pyri, co-infection, bacterial-fungal interactions, pathogenicity

陈春桥, 孙丽英, 盛强, 韩剑, 傅本重, 罗明. 梨火疫病菌与梨腐烂病菌对生长和致病的相互影响[J]. 园艺学报, 2024, 51(11): 2668-2684.

CHEN Chunqiao, SUN Liying, SHENG Qiang, HAN Jian, FU Benzhong, LUO Ming. Influences of Interactions Between Erwinia amylovora and Valsa pyri on Growth and Pathogenicity[J]. Acta Horticulturae Sinica, 2024, 51(11): 2668-2684.

图/表 14

表1 梨火疫病菌 hrp基因的荧光定量引物序列

Table 1 Sequence of fluorescent quantitative primers for the hrp genes of Erwinia amylovora

基因名称 Gene name	上游引物序列（5′-3′） Forward primer	下游引物序列（5′-3′） Reverse primer
HrpA	TGGTATCGCAGGCAGTTC	TCAGTATAGCTTTAGCGC
hrpB	GCGCGGAATAATGACGTC	TATCCAGACTCAGTACGC
hrpD	CAACTGAATCAACATCGG	GAAAATACTGACTGTTTC
hrpJ	CTCTGCATGACTTTGCCGAACA	TCAGGTTTAGCAGACGGCGCGTT
hrcJ	TGAGCATCGCGAGAAGAA	GAACAGTAGACGAAGCCA
HrpW	AATAGGCACAGCAGGAAC	CGCTCAGGTTAGTATCGG
hrpY	GATGTCATCATGCGTGAGGGACT	CGATTAAGCGTGCATTATGGGTAC
16S rRNA	AGAGTTAGATCATGGCTCAG	ACGGTTACCTAGTTACGACTT

表2 梨腐烂病菌细胞壁降解酶基因的荧光定量引物序列

Table 2 Sequences of fluorescent quantitative primers for cell wall degrading enzyme genes of Valsa pyri

基因名称 Gene name	上游引物序列（5′-3′） Forward primer	下游引物序列（5′-3′） Reverse primer
PG	CTATCTGCACCAGAGGACA	ATCTTCACGACCAAGTAATT
β-Gal	ATGAACGGAAAGTCCCCAC	TCCAATGAGCCATACACGG
PME	GATGCCTTGGAGTGGAGA	TGCTAATGTATAGCGTTC
α-Af	AGATACGCCTATCCTGAC	CACGGCATACTCGCTCAC
18S rRNA	CCATTGGAGGGCAAGTCT	GGTTCTCACGCTACACGA

图1 梨火疫病菌E.a001与梨腐烂病菌V.pKRL、梨黑斑病菌A.a发酵滤液共培养的E.a001生长曲线同一时间点不同字母表示经邓肯氏新复极差法检验在P < 0.05水平差异显著。下同。

Fig. 1 Growth curves of E.a001 co-cultured with V.pKRL，A.a fermentation filtrate Different letters at the same time point indicated significant difference at the P < 0.05 level by Duncan’s new multiple range test. The same below.

图2 梨火疫病菌E.a001与梨腐烂病菌V.pKRL、梨黑斑病菌A.a发酵滤液共培养液的pH值图中数据为平均数 ± 标准差，不同字母表示经邓肯氏新复极差法检验在P < 0.05水平差异显著。下同。

Fig. 2 pH values of co-cultures of E. a001 with V.pKRL，A.a fermentation filtrate Data are mean ± SD，different letters indicate significant difference at P < 0.05 level by Duncan’s multiple range test. The same below.

图3 梨腐烂病菌V.pKRL与梨火疫病菌E.a001、梨花枯病菌P.s、梨锈水病菌D.f发酵滤液共培养的V.pKRL生长曲线

Fig. 3 Growth curves of V.pKRL co-cultured with E.a001，P.s，D.f fermentation filtrates

图4 梨腐烂病菌V.pKRL与梨火疫病菌E.a001、梨花枯病菌P.s、梨锈水病菌D.f发酵滤液共培养液的pH值

Fig. 4 pH values of co-cultures of V.pKRL with E.a001，P.s and D.f fermentation filtrate

图5 梨火疫病菌E.a001与梨腐烂病菌V.pKRL发酵滤液共培养对E.a001的hrp基因表达量的影响

Fig. 5 Effects of E.a001 and V.pKRL fermentation filtrate co-culture on hrp gene expression in E.a001

图6 梨腐烂病菌V.pKRL与梨火疫病菌E.a001发酵滤液共培养对V.pKRL主要致病基因表达量的影响

Fig. 6 Effects of V.pKRL and E.a001 fermentation filtrates on of major pathogenic genes expression in V.pKRL

图7 香梨枝条上接种梨火疫病菌E.a001和梨腐烂病菌V.pKRL的不同方式 A：对照；B：E.a001单菌接种；C：V.pKRL单菌接种；D：E.a001（先）/V.pKRL（后）同位点依次复合接种；E：V.pKRL（先）/E.a001（后）同位点依次复合接种；F：E.a001 + V.pKRL异位点同时复合接种。

Fig. 7 The different inoculations methods of E. a001 and V. pKRL on Korla fragrant pear branches A：The control；B：E.a001 inoculation alone；C：V.pKRL inoculation alone；D：E.a001（first）/V.pKRL（next）sequential mixed inoculation at same site；E：V.p KRL（first）/E.a001（next）sequential mixed inoculation at same site；F：E.a001+ V.pKRL mixed inoculation at different sites.

表3 不同方式接种梨火疫病菌E.a001和梨腐烂病菌V.pKRL在香梨枝条上的致病力比较

Table 3 Comparison of pathogenicity by different inoculations with E. a001 and V. pKRL on pear branches

接种方式 Inoculation method		显症时间/d Time of appearance symptoms	病斑长度/cm Length of necrotic lesions			病斑面积/cm² Area of necrotic lesions	产孢数/ （个 · cm^-2） Spore number
接种方式 Inoculation method		显症时间/d Time of appearance symptoms	长度/cm Length	扩展/ （cm · d^-1） Expansion	病斑占枝条的比例/% Necrosis length/ length of the branches	病斑面积/cm² Area of necrotic lesions	产孢数/ （个 · cm^-2） Spore number
E.a001		3	10.38 ± 0.56 b	2.07 ± 0.12 b	48.00 ± 0.06 b	4.27 ± 0.89 b	—
V.pKRL		4	11.03 ± 0.77 b	2.21 ± 0.2 5 b	52.00 ± 0.13 b	3.31 ± 1.12 b	13
E.a001（先）/V.pKRL（后）同位点双菌 E.a001（first）/V.pKRL（next）at same site		2	13.06 ± 0.86 a	2.61 ± 0.17 a	58.00 ± 0.21 a	7.45 ± 0.73 a	22
V.pKRL（先）/E.a001（后）同位点双菌 V.pKRL（first）/E.a001（next）at same site		2	12.25 ± 0.66 a	2.42 ± 0.09 a	56.00 ± 0.14 a	7.13 ± 1.02 a	29
E.a001 + V.pKRL异位点双菌 E.a001 + V.pKRL at different sites	E. a001	3	9.75 ± 0.74 c	1.95 ± 0.33 c	42.00 ± 0.04 c	3.96 ± 0.94 c	—
	V. pKRL	3	11.16 ± 0.72 b	2.23 ± 0.18 b	50.00 ± 0.09 b	4.03 ± 0.65 b	17

图8 不同方式接种梨火疫病菌E.a001和梨腐烂病菌V.pKRL在香梨叶片上的症状 A：对照；B：E.a001单菌接种；C：V.pKRL单菌接种；D：E.a001（先）/V.pKRL（后）同位点依次复合接种；E：V.pKRL（先）/E.a001（后）同位点依次复合接种；F：E.a001 + V.pKRL异位点同时复合接种。

Fig. 8 Pathogenicity of different inoculations of E.a001 and V.pKRL on pear leaves A：The control；B：E.a001 inoculation alone；C：V.pKRL inoculation alone；D：Ea001（first）V.pKRL（next）sequential mixed inoculation at same site；E：V.pKRL（first）/E.a001（next）sequential mixed inoculation at same site； F：E.a001+ V.pKRL mixed inoculation at different sites.

图9 不同方式接种梨火疫病菌E.a001和梨腐烂病菌V.pKRL在香梨幼果上的症状 A：对照；B：E.a001单菌接种；C：V.pKRL单菌接种；D：E.a001（先）/V.pKRL（后）同位点先后复合接种；E：V.pKRL（先）/E.a001（后）同位点先后复合接种；F：E.a001 + V.pKRL异位点同时复合接种。

Fig. 9 Pathogenicity of different inoculations of E.a001 and V.pKRL on pear young fruits A：The control；B：E.a001 inoculation alone；C：V.pKRL inoculation alone；D：E.a001（first）/V.pKRL（next）sequential mixed inoculation at same site；E：V.pKRL（first）/E.a001（next）sequential mixed inoculation at same site；F：E.a001 + V.pKRL mixed inoculation at different sites.

表4 不同方式接种梨火疫病菌E.a001和梨腐烂病菌V.pKRL在香梨叶片上的致病力比较

Table 4 Comparison of pathogenicity by different inoculations with E.a001 and V.pKRL on pear leaves

接种方式 Inoculation method		显症时间/d Time of appearance symptoms	病斑长度/cm length of necrotic lesions		病斑面积/cm² Area of necrotic lesions	产孢数/ （个 · cm^-2） Spore number
接种方式 Inoculation method		显症时间/d Time of appearance symptoms	长度/cm Length	扩展/（cm · d^-1） Expansion	病斑面积/cm² Area of necrotic lesions	产孢数/ （个 · cm^-2） Spore number
E.a001		3	3.55 ± 1.18 c	0.71 ± 0.05 c	8.49 ± 1.09 b	—
V.pKRL		3	4.26 ± 1.41 b	0.85 ± 0.11 b	10.16 ± 1.52 b	9
E.a001（先）/V.pKRL（后）同位点双菌 E.a001（first）/V.pKRL（next）at same site		2	5.04 ± 1.76 a	1.01 ± 0.07 a	14.71 ± 0.78 a	19
V.pKRL（先）/E.a001（后）同位点双菌 V.pKRL（first）/E.a001（next）at same site		2	4.87 ± 1.08 a	0.97 ± 0.13 a	14.25 ± 1.22 a	17
E.a001 + V.pKRL 异位点双菌 E.a001 + V.pKRL at different sites	E.a001	3	3.46 ± 1.13 c	0.69 ± 0.08 c	7.97 ± 0.31 c	—
	V.pKRL	3	4.18 ± 1.73 b	0.83 ± 0.06 b	10.84 ± 0.37 b	11

表5 不同方式接种梨火疫病菌E.a001和梨腐烂病菌V.pKRL在香梨幼果上的致病力比较

Table 5 Comparison of pathogenicity by different inoculations with E.a001 and V.pKRL on pear young fruits

接种方式 Inoculation method		显症时间/d Time of appearance symptoms	病斑直径/cm Diameter of necrotic lesions		病斑面积/cm² Area of necrotic lesions	产孢数/ （个 · cm^-2） Spore number
接种方式 Inoculation method		显症时间/d Time of appearance symptoms	直径/cm Diameter	扩展/（cm · d^-1） Expansion	病斑面积/cm² Area of necrotic lesions	产孢数/ （个 · cm^-2） Spore number
E.a001		2	1.67 ± 0.30 c	0.33 ± 0.04 b	2.03 ± 0.18 c	—
V.pKRL		2	1.85 ± 0.31 b	0.37 ± 0.05 b	2.51 ± 0.19 b	18
E.a001（先）/V.pKRL（后）同位点双菌 E.a001（first）/V.pKRL（next）at same site		1	2.29 ± 0.34 a	0.46 ± 0.04 a	3.87 ± 0.19 a	26
V.pKRL（先）/E.a001（后）同位点双菌 V.pKRL（first）/E.a001（next）at same site		1	2.18 ± 0.24 a	0.43 ± 0.08 a	3.82 ± 0.13 a	23
E.a001 + V.pKRL异位点双菌 E.a001 + V.pKRL at different sites	E.a001	2	1.73 ± 0.42 c	0.35 ± 0.03 b	2.12 ± 0.23 c	—
	V.pKRL	2	1.78 ± 0.51 c	0.36 ± 0.05 b	2.19 ± 0.14 c	14

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梨火疫病菌与梨腐烂病菌对生长和致病的相互影响

Influences of Interactions Between Erwinia amylovora and Valsa pyri on Growth and Pathogenicity

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梨火疫病菌与梨腐烂病菌对生长和致病的相互影响

Influences of Interactions Between Erwinia amylovora and Valsa pyri on Growth and Pathogenicity

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