CN113621626B - FoCcw14基因在调控内生真菌-水稻共生体系抗稻瘟病中的用途 - Google Patents
FoCcw14基因在调控内生真菌-水稻共生体系抗稻瘟病中的用途 Download PDFInfo
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- C12N15/902—Stable introduction of foreign DNA into chromosome using homologous recombination
Abstract
本发明公开了FoCcw14基因在调控瓶霉属内生真菌‑水稻共生体系抗稻瘟病中的用途，属于植物病害防治技术领域。所述FoCcw14基因的核苷酸序列如SEQ ID NO.1所示。本发明通过对FoCcw14基因的克隆和分析，并利用同源重组的技术对该基因进行功能验证，发现敲除FoCcw14基因导致内生真菌‑水稻共生体系抗稻瘟病的性能显著降低。本发明为后续水稻育种提供了新的思路。
Description
技术领域
本发明涉及植物病害防治技术领域，具体涉及FoCcw14基因在调控瓶霉属内生真菌-水稻共生体系抗稻瘟病中的用途。
背景技术
稻瘟病是水稻重要病害之一，可引起大幅度减产，严重时减产 40％～50％，甚至颗粒无收。该病为通过气流传播的流行病，对水稻生产威胁极大，危害程度因品种、栽培技术以及气候条件不同有差别，世界各稻区均有发生，其中以叶部、节部发生为多，发生后可造成不同程度减产，尤其穗颈瘟或节瘟发生早而重，可造成白穗以致绝产。
生物防治是植物病害防控的研究热点，生物防治是利用有益微生物对病原菌产生各种不利作用(例如，抗菌，溶菌、竞争、重寄生等)，降低病原菌的数量或致病性；同时，用于生物防治的有益微生物还能诱导植物抗病性增强，提高植物免疫力，延缓、减轻或者抑制病害的发生。
其中植物内生真菌就是自然界存在的有益微生物之一，内生真菌至少有一部分生活史在健康的植物组织内部，定殖并吸取寄主内部的营养，但是与致病真菌不同的是，内生真菌并不会对寄主产生明显的伤害，可以在寄主的内部与其保持平衡的关系。
内生真菌在与植物互作过程中，展现出了许多有益的生物关系，例如促进宿主的生长，增加宿主的产量；产生毒素保护植物免受昆虫、食草动物的取食；在植物面临非生物胁迫逆境如高低温、重金属胁迫时，可以调控植物的生理生化反应，从而帮助其渡过不良环境；此外，内生真菌的定殖还可以诱导宿主产生系统抗病性，从而抵御真菌、细菌、线虫、病毒等的侵害。
在抗稻瘟病害胁迫方面，申请号为201911150894.1的专利文献公开甘蔗内生解淀粉芽孢杆菌CGB15菌株对甘蔗鞭黑粉菌、稻瘟菌、荔枝霜疫酶菌、荔枝炭疽菌、香蕉枯萎病菌都具有强烈的抑制作用。申请号为201610064924.7的专利文献公开从忽地笑鳞茎中分离的拮抗细菌菌株 HDXY对稻瘟病菌具有十分明显的拮抗作用。目前尚无报道指出野生稻内生真菌菌株在防治水稻穗颈瘟方面的功能。
目前对于内生真菌如何诱导宿主抗稻瘟病害胁迫的分子机制研究甚少，因此，通过探究内生真菌互作的分子机理，挖掘相关基因，并利用分子生物学手段改善内生真菌相关性能，以提升互作对象的抗病性，将其开发成生物防控制剂应用于农业生产，前景广阔。
发明内容
本发明的目的在于提供一种从水稻内生真菌中克隆得到的具有促进宿主水稻抗稻瘟病害胁迫特性的基因，为内生真菌-水稻共生体系的改良与生产提供了理论依据和相关基因。
为实现上述目的，本发明从保藏号为CCTCC NO:M 2021505的瓶霉属内生真菌Falciphora oryzae FO-R20中鉴定克隆到具有促进宿主水稻抗稻瘟病害胁迫特性的基因FoCcw14，该基因的核苷酸序列如SEQ ID NO.1 所示。该基因CDS区全长453bp，编码一个150aa的蛋白序列。所述 FoCcw14基因编码的蛋白质的氨基酸序列如SEQ ID No.2所示。
本发明利用同源重组的技术，对内生真菌Falciphora oryzae FO-R20 基因组中的FoCcw14基因进行敲除，获得突变体ΔFoCcw14，发现相较于内生真菌野生型-水稻共生体系，该突变体在与水稻根部共培养以后，水稻的系统抗稻瘟病害的性能显著降低了15％，该结果表明，FoCcw14基因是调控内生真菌影响宿主抗病性的关键基因。
因此，本发明提供了核苷酸序列如SEQ ID NO.1所示的FoCcw14基因在调控瓶霉属内生真菌-水稻共生体系抗稻瘟病中的用途。该基因增强共生系统对稻瘟病菌病害的抗性。FoCcw14基因敲除后使得瓶霉属内生真菌-水稻共生体系抗稻瘟病菌病害的功能降低。
进一步地，所述瓶霉属内生真菌-水稻共生体系的构建方法包括：将内生真菌Falciphora oryzae FO-R20与水稻共培养，使其定殖于水稻根部组织中。
优选的，共培养前，将内生真菌Falciphora oryzae FO-R20菌株接种于PDA培养基进行活化培养，25℃黑暗培养7天。
优选的，所述共培养包括将萌发后的水稻种子与内生真菌Falciphora oryzaeFO-R20菌丝块接种于无菌的1/2MS培养基上进行培养。
优选的，共培养的条件为：22-25℃下培养20-25天，每天光照16h，暗培养8h。
本发明还提供了一种防治水稻稻瘟病的方法，包括：将核苷酸序列如 SEQ IDNO.1所示的FoCcw14基因导入瓶霉属内生真菌获得重组菌，再将重组菌与水稻共培养，获得重组菌-水稻共生体系。
本发明具备的有益效果：
本发明首次公开水稻内生真菌中影响宿主抗稻瘟病性能的基因 FoCcw14，通过对FoCcw14基因的克隆和分析，并利用同源重组的技术对该基因进行功能验证，发现敲除FoCcw14基因导致内生真菌-水稻共生体系抗稻瘟病的性能显著降低。本发明为后续水稻育种提供了新的思路。
附图说明
图1为内生真菌Falciphora oryzae FO-R20中FoCcw14的生物信息学分析，图A为多序列联配，图B为SMART预测结果，图C为系统发育分析。
图2为内生真菌基因突变体ΔFoCcw14菌株。
图3为内生真菌基因突变体ΔFoCcw14菌株在水稻根部的定殖后，水稻根部横切面，左图为白光下的视图，右图为488nm激光下的视图。
图4为内生真菌基因突变体ΔFoCcw14菌株在水稻根部的定殖后并喷施稻瘟病菌，发病时病斑面积。
具体实施方式
下面通过具体实施例，对本发明的技术方案作进一步的具体说明。应当理解，本发明的实施并不局限于下面的实施例，对本发明所做的任何形式上的变通和/或改变都将落入本发明保护范围。
在本发明中，若非特指，所采用的设备和原料等均可从市场购得或是本领域常用的。下述实施例中的方法，如无特别说明，均为本领域的常规方法。
本发明以课题组前期工作从云南疣粒野生稻根系中分离得出一株新的瓶霉属内生真菌Falciphora oryzae FO-R20为材料，敲除FoCcw14基因，对阐明内生真菌FO-R20-水稻共生体系响应稻瘟病菌胁迫的分子机理具有重要意义。
实施例1 FoCcw14基因分析
内生真菌Falciphora oryzae FO-R20分离自疣粒野生稻根系，2021年 5月8日保藏于中国.武汉.武汉大学的中国典型培养物保藏中心，保藏号为：CCTCC NO:M 2021505。
前期研究表明，内生真菌Falciphora oryzae FO-R20定殖于水稻植株根部组织中，可以显著提升水稻对稻瘟病害胁迫的抗性。
对Falciphora oryzae FO-R20进行全基因组序列分析，通过在线软件 SMART预测蛋白质的信号肽、结构域。通过多序列联配和系统发育分析，确定FoCcw14为研究对象。FoCcw14基因的核苷酸序列如SEQ ID NO.1 所示，其编码的氨基酸序列如SEQ ID NO.2所示，其生物信息学分析结果如图1所示。
实施例2验证FoCcw14基因功能
供试内生真菌：Falciphora oryzae FO-R20；
供试植物：水稻Oryza sativa L.，常规品种，CO39。
1、菌株培养
将保存于滤纸片上的FO-R20菌株接种于马铃薯葡萄糖琼脂(PDA)固体培养基上进行活化培养，25℃，黑暗培养7d，备用。
PDA培养基：每升含葡萄糖20g，马铃薯200g，琼脂15g。根据待配培养基的体积称取所需马铃薯，水煮后捣碎溶解过滤，加入葡萄糖和琼脂，121℃高压蒸汽灭菌20min。
2、基因突变体菌株ΔFoCcw14的获得
设计目的基因上下游引物，扩增上下游片段和SUR片段。具体引物序列如下：
FoCcw14-UP-F：
5’-GGTACCCGGGGATCCTCTAGATTACCTGCCTACCTACCTAG-3’；
FoCcw14-UP-R：
5’-GGCACTGTGGCGTTGGCACGCGTCTGCTGTGTCTTGAC-3’；
FoCcw14-DOWN-F：
5’-GGGAATTGCATGCTCTCACTCATTCCTTCTAGTGCTTCA-3’；
FoCcw14-DOWN-R：
5’-ACGACGGCCAGTGCCAAGCTTGAATACTACAAGAACCGTGAC-3’；
SUR-F：5’-GTGCCAACGCCACAGTGCCCCAC-3’；
SUR-R：5’-GATTGTGAATCGTGAGAGCATGCAATTC-3’。
通过限制性内切酶XbaI和HindⅢ酶切载体pKO1B。根据ClonExpress II One Stepclone Kit(Vazyme，中国)的操作说明，将回收后的片段和酶切后的pKO1B载体连接在一起，构建基因敲除载体并在大肠杆菌感受态DH5α中进行转化。挑出长好的转化子放入1mL LB液体培养基 (含50μg/ml卡那霉素)中，37℃，200rpm震荡4-6h，吸取1-2μL菌液加入到验证体系中，进行PCR扩增和凝胶电泳。
根据AxyPrep-96(Axygen，US)的操作指南进行质粒的提取。并将质粒转入农杆菌感受态AGL1中，通过农杆菌介导的转化方法获得突变体转化子，提取转化子的DNA进行长短片段验证和拷贝数验证，将验证正确的转化子接种到PDA培养基上进行后续实验，并接种到冻存管的PDA 斜面上进行保存。
突变体菌株ΔFoCcw14的菌落形态如图2所示，通过实验发现，突变体菌株ΔFoCcw14在菌落形态和直径上与野生型基本一致。
3、FO-R20野生型菌株、突变体菌株ΔFoCcw14与水稻根部共培养
将水稻种子去壳后放置在三角瓶中，用1％的NaClO表面消毒15min，用无菌水清洗3次后备用。将消毒后的种子均匀平铺在1/2MS(Murashige and Skoog)培养基上，用Parafilm封口膜封口，置于25℃植物培养箱(16h光照/8h黑暗)。3天后种子露白，将其移入含1/2MS培养基的组培瓶中，每瓶10颗种子。同时接入3块菌饼(直径为0.5cm)。对照组为无菌PDA琼脂块。设3个重复。
如图3所示，突变体GFP标记的菌丝只在表皮层扩展，无法进入到水稻细胞内部。
4、稻瘟病菌分生孢子的喷施
将稻瘟病菌菌株Guy11接种于CM固体培养基中，25℃光照培养10-12 天。用无菌水将Guy11分生孢子洗下，三层滤纸过滤，配置成孢子悬浮液，浓度为1×105。随后，配置0.4％明胶溶液，按体积比1:1与孢子悬浮液混合，使稻瘟病菌孢子悬浮液终浓度为5×104。用喷雾器将孢子悬浮液均匀地喷洒到水稻苗叶片上，每瓶用量1mL。然后，将组培瓶置于25℃植物培养箱，黑暗培养2天。2天后，补充光照，16h光照/8h黑暗，培养4-5 天，记录叶片发病情况。
CM培养基(1L)：Yeast Extract(1g)，Casamino acid(1g)，D-glucose (10g)，KH2PO4(1.52g)，NaNO3(6g)，Peptone140(2g)，KCl(0.52g)， MgSO4·7H2O(0.52g)，0.1％(v/v)Vitamin solution，0.1％(v/v)Trace Element。用NaOH调节PH至6.5，固体培养基中加入15g/L琼脂。采用高压蒸汽灭菌，条件为121℃，15min。
5、稻瘟病病斑率的统计
将发病的水稻叶片贴在白纸上，放上标尺拍照。在Photoshop中随机选取5cm长度叶片，统计所选叶片的全像素，调节容差，选取绿色无病斑面积，统计其像素。
病斑率(％)的计算公式为：(截取叶片全像素-绿色无病斑面积像素)/截取叶片的全像素。
如图4所示，空白对照和与突变体ΔFoCcw14共生的水稻叶片出现典型的纺锤状病斑。野生型菌株共生的水稻叶片上，病斑小且少。可见， FoCcw14基因在调控内生真菌Falciphora oryzae FO-R20-水稻共生体系响应稻瘟病菌胁迫时发挥重要作用。
序列表
<110> 浙江省农业科学院
<120> FoCcw14基因在调控内生真菌-水稻共生体系抗稻瘟病中的用途
<160> 8
<170> SIPOSequenceListing 1.0
<210> 1
<211> 453
<212> DNA
<213> Falciphora oryzae
<400> 1
atgaagacct ccgccgtcct cgcagccctg gcgctcgccg tcgccaccaa cgcgcagaca 60
gccggcgacc tgcccaagtg cgctcaagac tgcgcgaacc agttcctcct gaacggcatc 120
ggcaactgcg gccgcgaccc aaagtgcatc tgctccaaca agtccttcat cggagacatc 180
tcgtgctgcc tggccaagcc gggcggctgc agcgccgacg accagaagaa ggccatcgcc 240
tacgccgccc agctgtgcca ggccaacggc gtcaccgtcc ccgaccaggt cgtctgcaac 300
agcaacagca acaacggcac cgcggccacc tctggcgccg gcggcgccgc ggccaccacc 360
tcgtcgtcca gggcctgggc cccgcaccag accggcgccc ctgctgctgt cggcctcctc 420
ggcggcctgg ctgctgtcgc cgtgctgctg tga 453
<210> 2
<211> 150
<212> PRT
<213> Falciphora oryzae
<400> 2
Met Lys Thr Ser Ala Val Leu Ala Ala Leu Ala Leu Ala Val Ala Thr
1 5 10 15
Asn Ala Gln Thr Ala Gly Asp Leu Pro Lys Cys Ala Gln Asp Cys Ala
20 25 30
Asn Gln Phe Leu Leu Asn Gly Ile Gly Asn Cys Gly Arg Asp Pro Lys
35 40 45
Cys Ile Cys Ser Asn Lys Ser Phe Ile Gly Asp Ile Ser Cys Cys Leu
50 55 60
Ala Lys Pro Gly Gly Cys Ser Ala Asp Asp Gln Lys Lys Ala Ile Ala
65 70 75 80
Tyr Ala Ala Gln Leu Cys Gln Ala Asn Gly Val Thr Val Pro Asp Gln
85 90 95
Val Val Cys Asn Ser Asn Ser Asn Asn Gly Thr Ala Ala Thr Ser Gly
100 105 110
Ala Gly Gly Ala Ala Ala Thr Thr Ser Ser Ser Arg Ala Trp Ala Pro
115 120 125
His Gln Thr Gly Ala Pro Ala Ala Val Gly Leu Leu Gly Gly Leu Ala
130 135 140
Ala Val Ala Val Leu Leu
145 150
<210> 3
<211> 41
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 3
ggtacccggg gatcctctag attacctgcc tacctaccta g 41
<210> 4
<211> 38
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 4
ggcactgtgg cgttggcacg cgtctgctgt gtcttgac 38
<210> 5
<211> 39
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 5
gggaattgca tgctctcact cattccttct agtgcttca 39
<210> 6
<211> 42
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 6
acgacggcca gtgccaagct tgaatactac aagaaccgtg ac 42
<210> 7
<211> 23
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 7
gtgccaacgc cacagtgccc cac 23
<210> 8
<211> 28
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 8
gattgtgaat cgtgagagca tgcaattc 28
Claims (7)
1.核苷酸序列如SEQ ID NO.1所示的FoCcw14基因在调控瓶霉属内生真菌-水稻共生体系抗稻瘟病中的用途，其特征在于，所述瓶霉属内生真菌为保藏号为CCTCC NO:M 2021505的Falciphora oryzae FO-R20。
2.如权利要求1所述的用途，其特征在于，所述FoCcw14基因编码的蛋白质的氨基酸序列如SEQ ID No.2所示。
3.如权利要求1所述的用途，其特征在于，瓶霉属内生真菌FoCcw14基因敲除后使得瓶霉属内生真菌-水稻共生体系抗稻瘟病菌病害的性能降低。
4.如权利要求1所述的用途，其特征在于，所述瓶霉属内生真菌-水稻共生体系的构建方法包括：将内生真菌Falciphora oryzae FO-R20与水稻共培养，使其定殖于水稻根部组织中。
5.如权利要求4所述的用途，其特征在于，共培养前，将内生真菌Falciphora oryzaeFO-R20菌株接种于PDA培养基进行活化培养，25℃黑暗培养7天。
6.如权利要求4所述的用途，其特征在于，所述共培养包括将萌发的水稻种子与内生真菌Falciphora oryzae FO-R20菌块接种于无菌的1/2MS培养基上进行培养。
7.如权利要求6所述的用途，其特征在于，共培养的条件为：22-25℃下培养20-25天，每天光照16h，暗培养8h。
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