新疆农业科学, 2025, 62(6): 1337-1343 DOI: 10.6048/j.issn.1001-4330.2025.06.005

作物遗传育种·耕作栽培·生理生化

链霉菌HU2014诱导小麦抗性基因的表达

朱红霞,1, 胡林峰,2, 王琦琦1, 张军高1, 张少民1, 周小云1, 雷斌2, 张志东,3

1.新疆维吾尔自治区农业科学院农业资源与环境研究所/新疆作物化学调控工程技术研究中心,乌鲁木齐 830091

2.新疆农业科学院农药试制中心/新疆作物化学调控工程技术研究中心,乌鲁木齐 830091

3.新疆维吾尔自治区农业科学院微生物研究所,乌鲁木齐 830091

Expression analysis of disease resistance-related genes in wheat induced by Streptomyces sp. HU2014

ZHU Hongxia,1, HU Linfeng,2, WANG Qiqi1, ZHANG Jungao1, ZHANG Shaomin1, ZHOU Xiaoyun1, LEI Bin2, ZHANG Zhidong,3

1. Institute of Agricultural Resources and Environment / Xinjiang crop chemical control engineering technology research center, Xinjiang Academy of Agricultural Sciences, Urumqi 830091, China

2. Pesticide Trial Production Center, Xinjiang Academy of Agricultural Sciences/Xinjiang Crop Chemical Control Engineering Technology Research Center, Urumqi 830091, China

3. Institute of Applied Microbiology, Xinjiang Academy of Agricultural Sciences, Urumqi 830091, China

通讯作者: 胡林峰(1981-),男,湖北天门人,副研究员,研究方向为生物农药资源开发与利用,(E-mail)wood9818@sina.com;张志东(1977-),男,河南永城人,研究员,博士,研究方向为特殊环境微生物及益生菌资源的挖掘和利用,(E-mail)Zhangzheedong@qq.com

收稿日期: 2024-12-5  

基金资助: 新疆维吾尔自治区科技创新科技援疆项目(2024E02002)
新疆农业科学院稳定支持项目(xjnkywdzc-2024001-04)
新疆农业科学院稳定支持项目(xjnkywdzc-2024003-70)

Corresponding authors: HU Linfeng (1981-), male, from Tianmen, Hubei, associate researcher, research direction: biological pesticide resource development and utilization, (E-mail)wood9818@sina.com;ZHANG Zhidong (1977-), male, from Yongcheng, Henan, researcher, Ph.D., research direction: the exploration and utilization of microorganisms in special environments and probiotics, (E-mail)Zhangzheedong@qq.com

Received: 2024-12-5  

Fund supported: S & T Assisting Xinjiang Project of Xinjiang Uygur Autonomous Region(2024E02002)
The Stable Support Project to Agricultural Sci-Tech Renovation of Xinjiang Academy of Agricultural Sciences(xjnkywdzc-2024001-04)
The Stable Support Project to Agricultural Sci-Tech Renovation of Xinjiang Academy of Agricultural Sciences(xjnkywdzc-2024003-70)

作者简介 About authors

朱红霞(1982-),女,山西汾阳人,助理研究员,研究方向为微生物资源挖掘与利用,(E-mail)zhxhg105@163.com

摘要

【目的】研究链霉菌HU2014处理对小麦抗性基因表达的动态规律,探讨该菌株诱导寄主植物产生抗病性的可能机制。【方法】采用链霉菌HU2014无细胞发酵滤液对矮抗58(AK58)、百农307(BN307)和周麦22(ZM22)等3个小麦品种进行灌根处理,分别于灌根后2、6、12、24、48和96 h采集叶片,利用实时荧光定量PCR技术检测小麦叶片中相关抗性基因表达量的变化。【结果】 HU2014菌株无细胞发酵液能够诱导4个防卫反应基因、1个水杨酸途径基因和1个茉莉酸途径基因的表达。其中TaPR1基因在处理AK58品种6 h后,BN307品种24 h后和ZM22样品48 h后表达量最大;TaPR2基因的表达在处理AK58的6 h后,BN307的12 h后和ZM22的24 h后达到最大水平;TaPR3基因表达量在AK58的2 h后,BN307的6 h后和ZM22的2 h后达到峰值;TaPR5基因在AK58的72 h后、BN307的96 h后和ZM22的12 h后达到峰值;PAL基因在AK58的48 h后、BN307的24 h后和ZM22的24 h后显示出最高的倍数变化;LOX基因表达在AK58、BN307和ZM22的6 h后达到峰值。【结论】TaPR1基因在AK58和ZM22中下调,TaPR3基因在AK58和BN307中下调。ZM22中TaPR2、TaPR3、TaPR5、PAL和LOX的最高表达水平高于AK58和BN307,而BN307中TaPR1的最高表达水平高于AK58和ZM22。链霉菌HU2014可以诱导小麦系统性获得抗性和诱导性系统抗性。

关键词： 链霉菌; 小麦; 抗病相关基因; 生物防治; 诱导抗病性

Abstract

【Objective】 The purpose of this study is to reveal the dynamic changes in the expression of wheat resistance genes under the treatment of Streptomyces sp. HU2014 and to explore the possible mechanism by which this strain induces disease resistance in the host. 【Methods】 In this paper, the cell-free fermentation filtrate of this strain was used to irrigate the roots of three wheat varieties Aikang 58 (AK58), Bainong 307 (BN307), and Zhoumai 22 (ZM22). After irrigation, leaves were collected at 2 h, 6 h, 12 h, 24 h, 48 h, and 96 h, and the changes in the expression of related resistance genes were detected by real-time fluorescent quantitative PCR technology. 【Results】 The fermentation filtrate of this strain could induce the expression of 4 defense response genes, 1 salicylic acid pathway gene, and 1 jasmonic acid pathway gene. Among them, the transcript of TaPR1 gene accumulated at highest levels at 6 h in AK58, 24 h in BN307, and 48 h in ZM22 respectively. The expression of TaPR2 gene reached the highest level at 6 hours in AK58, 12 h in BN307, and 24 h in ZM22 respectively. The transcript of TaPR3 gene reached its peak at 2 h in AK58, 6 h in BN307, and 2 h in ZM22 respectively. TaPR5 gene reached its peak at 72 h in AK58, 96 h in BN307, and 12 h in ZM22 respectively. The PAL gene showed the highest fold change at 48 h in AK58, 24 h in BN307, and 24 h in ZM22 respectively. The expression of LOX gene reached its peak at 6 h in AK58, BN307, and ZM22 respectively. 【Conclusion】 The results indicate that the TaPR1 gene is down-regulated in AK58 and ZM22, and the TaPR3 gene is down-regulated in AK58 and BN307. The highest expression levels of TaPR2, TaPR3, TaPR5, PAL, and LOX in ZM22 are higher than those in AK58 and BN307, while the highest expression level of TaPR1 in BN307 is higher than that in AK58 and ZM22. Streptomyces sp. HU2014 can induce systemic acquired resistance and systemic resistance in wheat.

Keywords： streptomyces; wheat; disease resistance related genes; biological control; inducing resistance

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本文引用格式

朱红霞, 胡林峰, 王琦琦, 张军高, 张少民, 周小云, 雷斌, 张志东. 链霉菌HU2014诱导小麦抗性基因的表达[J]. 新疆农业科学, 2025, 62(6): 1337-1343 DOI:10.6048/j.issn.1001-4330.2025.06.005

ZHU Hongxia, HU Linfeng, WANG Qiqi, ZHANG Jungao, ZHANG Shaomin, ZHOU Xiaoyun, LEI Bin, ZHANG Zhidong. Expression analysis of disease resistance-related genes in wheat induced by Streptomyces sp. HU2014[J]. Xinjiang Agricultural Sciences, 2025, 62(6): 1337-1343 DOI:10.6048/j.issn.1001-4330.2025.06.005

0 引言

【研究意义】我国小麦栽培面积和总产量均居谷物第一位,但禾谷丝核菌引起的小麦纹枯病则导致小麦不同程度减产。特别是随着秸秆粉碎入田耕作制度的实行,小麦纹枯病的发病率逐年提高[1]。截至目前尚无对小麦纹枯病菌具有有效抗性的小麦品种[2]。与不同的防治策略相比,利用环境友好的微生物进行生物防治被认为是防治小麦纹枯病的一种有效途径[3]。【前人研究进展】作为最大的放线菌属,链霉菌占土壤放线菌总数的50%[4]。这些链霉菌产生丰富多样的次级代谢产物,不仅可以促进植物生长,还具有对多种植物病原菌的生物防治潜力[5-9]。链霉菌具有生防潜力的机理之一就是其能诱导植物的防御机制,增强植物的抗病性[10,11]。病程蛋白(PR蛋白)是植物与病原物的接触识别以及抗病性产生过程中的产物,其能够抵抗病原物的再次侵染,并在体外表现出一定的抗菌作用,在植物抗病性中起重要作用[12-15]。例如,β-1,3-葡聚糖酶(PR2)可以抑制病原菌的生长,其主要通过破坏病原菌细胞壁并从中释放激发因子,诱导植株产生抵抗病原菌的系统抗性[16];苯丙氨酸解氨酶(PAL)是植物体内次级代谢反应的关键酶和限速酶,在植物抵御病虫害中发挥着独特的作用[17];脂氧合酶(LOX)在植物体中可能参与植物的生长发育、成熟、衰老和防御过程的调控[18-20]。【本研究切入点】实时荧光定量PCR技术是一项用来检测和研究目的基因的表达量和动态变化的常用技术,具有很强的灵敏度和准确性,可以同时实现定性和定量的分析,是开展作物抗逆研究的重要手段[21]。前期研究表明选用链霉菌HU2014及其无细胞发酵液在体外可有效抑制小麦纹枯病菌的生长,需研究链霉菌HU2014处理对小麦抗性基因表达的动态规律。【拟解决的关键问题】采用HU2014无细胞发酵液对小麦进行灌根处理,采用实时荧光定量PCR技术对小麦相关抗病基因在不同时段的表达量进行检测,从诱导抗病方面揭示该菌株及其代谢产物对小麦抗病能力的作用机制。

1 材料与方法

1.1 材料

试验在新疆维吾尔自治区农业科学院新疆作物化学调控工程技术研究中心温室进行。3个小麦品种矮抗58、百农307和周麦22由新疆农业科学院新疆作物化学调控工程技术研究中心提供。

葡萄糖大豆蛋白胨酵母提取物培养基(GPY):葡萄糖20 g,大豆蛋白胨5 g,酵母提取物5 g,去离子水1 000 mL。

马铃薯葡萄糖琼脂培养基(PDA):马铃薯200 g,葡萄糖20 g,琼脂粉15 g,去离子水1 000 mL。

1.2 方法

1.2.1 链霉菌HU2014发酵液的制备

保存的链霉菌HU2014在PDA固体培养基上活化后,用打孔器将菌株打成直径5 mm的菌饼,将其接种到GPY液体培养基里,于25℃、150 r/min下摇床培养12 d。将得到的发酵原液先用筛绢初滤,再用0.25 mm针头滤芯过滤得无细胞发酵液,稀释500倍后对小麦进行灌根处理。

1.2.2 试验设计

将3个供试小麦品种催芽后播种在花盆中(每盆15株,12 cm×12 cm×9 cm),于温室中培养(25℃ / 20℃)±1℃,光照/黑暗:14 h /10 h,相对湿度65%左右,正常浇灌。待长出2叶1心的小麦苗后,按照每盆100 mL进行灌根处理。灌根前为0 h,灌根后2、6、12、24、48和96 h,分别采集100 mg叶片放入1.5 mL离心管中,立刻投入液氮中冷冻3~5 min,再放置于-80℃超低温冰箱中保存。

1.2.3 RNA提取,cDNA合成及实时荧光定量PCR

采用Trizol法提取小麦叶片总RNA。参照Prime ScriptTM RT reagent kit with gDNA Eraser(TaKaRa,中国,大连)中说明书的方法合成第一链cDNA。以1 mg总RNA为模板,RT Primer Mix为引物,在200 mL的RNase-Free离心管中进行反应。以Actin作为内参基因,通过Applied biosystems 7500 Real-Time PCR System,检测各个设定基因的转录量,比较小麦经供试药剂诱导后的基因表达量差异,反应程序:95℃ 30 s; 95℃ 5 s; 60℃ 34s,40个循环。试验重复3次,结果以小麦目标基因与内参基因(Actin)的拷贝数比值表示。采用2-ΔΔCt方法进行相对表达量的计算分析,采用SPSS18软件(IBM,美国,纽约)进行显著性分析。定量PCR引物设计参照Genbank数据库。表1

表1   定量PCR引物序列

Tab.1  Quantitative PCR primer sequence used in this study

引物
Gene name		上下游引物
Forward and reverse primers (5'-3')
TaActinGGACCTCACGGATAATCTAATG
TGACCATCAGGCATCTCA
TaPR1AACCTCGGCGTCTTCATCA
TTTACTCGCTCGGTCCCTCT
TaPR2TGCCGTTGCTCTCTTCAT
ATGCCCTTGGACCTGTAGA
TaPR3AGAGATAAGCAAGGCCACGTC
GGTTGCTCACCAGGTCCTTC
TaPR5ACAGCTACGCCAAGGACGAC
CGCGTCCTAATCTAAGGGCAG
PALCCAATGTTCTGTCCGTCCT
GCTGCTTCAATCTGTCCAG
LOXAGGCAACTACATCTACGCTTC
GCCATCAACACCAGAGTCA

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2 结果与分析

2.1 HU2014诱导AK58抗性基因表达

研究表明,在HU2014发酵液处理小麦品种AK58后,TaPR1基因表达量在随后的几个接种时间与对照相比,显著降低。TaPR2基因在6 h与对照相比显著升高,其余时间点均显著降低。TaPR3基因表达量与对照相比,除2 h与对照相比无显著差异外,其余时间点均表达显著降低。TaPR5基因的表达量呈现降低趋势(除72 h无显著差异外)。LOX基因在6 h与对照相比显著升高,其余时间点均显著降低。PAL基因的表达在48和96 h时与对照比无显著差异,其余时间点均表达显著降低。图1

图1

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图1   链霉菌HU2014对小麦AK58的抗性基因表达

Fig.1   The strain HU2014 on the expression of resistance genes in wheat (cultivar AK58) plant


2.2 HU2014诱导BN307抗性基因表达

研究表明,在HU2014发酵液处理小麦品种BN307后,6个基因均在不同时间点显著表达。TaPR1基因在24 h后的积累最高,与对照相比,倍数值为6.95。TaPR2基因的表达在24 h 时与对照相比达到最高水平,其倍数值为55.70,其余时间点均表达显著降低。TaPR3基因表达量在随后的几个时间点与对照相比均显著降低。TaPR5基因的表达量在24 h出现小高峰,此后急剧降低,但到96 h积累最高,其倍数值为3.33。LOX基因在6和24 h均表达量最高,其余时间点均显著降低。PAL基因的表达在24和48 h时积累最高,其余时间点均表达显著降低。图2

图2

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图2   链霉菌HU2014对小麦BN307的抗性基因表达

Fig.2   The strain HU2014 on the expression of resistance genes in wheat (cultivar BN307) plant


2.3 HU2014诱导ZM22抗性基因表达

研究表明,在HU2014发酵液处理小麦品种ZM22后,除TaPR1基因表达量在对照点时间后均显著降低外,其余5个基因均在不同的处理时间点显著升高。TaPR2基因的表达在12 h 时与对照相比达到最高水平,其倍数值为105.47,其余时间点均表达显著降低。TaPR3基因表达量在2 h最高,其余时间点均显著降低。TaPR5基因的表达量在6 h出现小高峰,此后急剧降低,但到12 h积累最高,其倍数值为98.67。LOX基因在6 h表达量最高,其余时间点均显著降低。PAL基因的表达在24和48 h积累较高,其余时间点均表达显著降低。图3

图3

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图3   链霉菌HU2014对小麦ZM22的抗性基因表达

Fig.3   The strain HU2014 on the expression of resistance genes in wheat (cultivar ZM22) plant


3 讨论

3.1 链霉菌除了通过产生拮抗化合物直接抑制病原菌外,还可以通过激活宿主植物的抗性途径间接为其植物宿主提供保护[22-24]。病程相关蛋白1(PR1)、β-1,3-葡聚糖酶(PR2)、几丁质酶(PR3)和类甜菜蛋白(PR5)是与防御相关的基因。研究利用荧光定量PCR技术对链霉菌HU2014处理小麦后,诱导小麦叶片相关抗性基因的表达谱进行表达分析。500倍稀释的HU2014发酵液在不同程度上触发了小麦的4个防御相关基因(TaPR1、TaPR2、TaPR3和TaPR5)的表达。PR1家族基因通常被用作疾病抗性的分子标记[25-27]。病原真菌细胞壁的主要成分是几丁质和β-1,3-葡聚糖,PR2酶水解真菌细胞壁外层的葡聚糖,PR3酶继而水解几丁质而发挥作用。因此,植物的PR2和PR3酶通过协同作用,有效抑制病原菌的繁殖和生长,在植物抗病途径中起着关键作用[28-31]。PR5在高等植物中广泛分布,其表达可以被各种生物和非生物胁迫诱导[32,33]。3个小麦品种的相关基因表达量在给定的所有试验点中,仅有一个点会出现表达最高峰,可能与小麦细胞内环境的动态变化和其分子层面的随机性有关。

3.2 苯丙氨酸解氨酶(PAL)是苯丙酸合成途径的限速酶,脂氧合酶(LOX)是茉莉酸途径的关键酶。水杨酸途径基因(PAL)和茉莉酸途径基因(LOX)在试验研究中也做了相应的表达分析。HU2014发酵液处理在不同程度上诱导了PAL和LOX酶的转录谱。该菌可能通过诱导小麦的水杨酸和(或)茉莉酸途径基因的表达,提高小麦的抗病性。相关研究也证明了这点,例如链霉菌N2中提取出来的抗真菌霉素N2处理水稻幼苗,PAL的表达量与对照组相比显著提高[34];在病原菌和不吸水链霉菌公主岭变种(农抗“769”)的双重诱导下,大豆中PAL的表达量比对照组提高了95.22%[35];经链霉菌JD211菌种的发酵液处理后的水稻叶片中LOX活性较对照组提高了40.85%[36]。

另外,从研究中可知,经HU2014无细胞发酵液处理后,TaPR1基因在AK58和ZM22中下调,TaPR3基因在AK58和BN307中下调。ZM22中TaPR2、TaPR3、TaPR5、PAL和LOX的最高表达水平高于AK58和BN307,而BN307中TaPR1的最高表达水平高于AK58和ZM22。以上结果表明同一抗病基因在不同小麦品种的表达谱中表现有差异,可能与小麦遗传背景有关。

试验基于HU2014的发酵液通过灌根的盆栽方式进行苗期小麦的抗性基因表达分析,而小麦在不同的生育阶段,基因表达存在差异,小麦的抗病性表现会有不同。因此,对小麦整个生育期的抗病性能还有待进一步研究。其次,该菌的发酵液对小麦根际土壤的影响也需要进行相关的研究,以揭示其诱导抗病性的相关机制。

4 结论

链霉菌HU2014诱导小麦抗性基因TaPR1、TaPR2、TaPR3、TaPR5、PAL和LOX的表达。TaPR1基因在AK58和ZM22中下调,TaPR3基因在AK58和BN307中下调。ZM22中TaPR2、TaPR3、TaPR5、PAL和LOX的最高表达水平高于AK58和BN307,而BN307中TaPR1的最高表达水平高于AK58和ZM22。链霉菌HU2014可以诱导小麦系统性获得抗性和诱导性系统抗性,并且这种诱导抗性作用可能与水杨酸和茉莉酸合成途径有关。

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