尖孢镰刀菌亚麻专化型<i>FolSid1</i>基因敲除及功能分析

doi:10.7668/hbnxb.20194757

摘要/Abstract

摘要：

通过克隆尖孢镰刀菌亚麻专化型FolSid1基因,确定该基因在镰刀菌中的生物学功能以及蛋白定位。通过与尖孢镰刀菌近缘菌同源比对,克隆得到FolSid1的基因序列,基于同源重组原理,使用Split Marker法构建含有潮霉素抗性基因(hph)的基因缺失盒,通过PEG介导法转入野生型原生质体中,获得基因敲除突变体(ΔFolSid1),对突变菌株做表型鉴定和致病力分析,分析该基因在尖孢镰刀菌亚麻专化型中的功能。构建含有FolSid1基因的绿色荧光表达载体pZESH1,对FolSid1-EGFP融合蛋白进行亚细胞定位。结果表明,FolSid1基因序列全长5 392 bp,含有3个内含子。与野生型和外插入突变体相比,尽管敲除突变体ΔFolSid1分生孢子形态和大小没有不同,但产量显著下降;形态学观察发现,敲除突变体的生长速度明显减慢;亚麻试验显示,缺失突变体的致病力明显降低;基因的亚细胞定位试验显示,FolSid1蛋白主要定位于菌丝细胞的细胞膜上。FolSid1基因能够调控尖孢镰刀菌亚麻专化型的菌丝营养生长、分生孢子发生且对亚麻具有致病性。

关键词: 尖孢镰刀菌亚麻专化型, FolSid1, Split Marker, 基因敲除

Abstract:

The aim was to study the biological function of FolSid1 gene in Fusarium oxysporum f.sp.lini and its protein localization in Fusarium by cloning the gene.The gene sequence of FolSid1 was cloned by homologous comparison with F.oxysporum, and based on the principle of homologous recombination, a gene deletion box containing hydromycin resistance gene(hph)was constructed by Split Marker strategy,and the gene deletion mutant(ΔFolSid1)was obtained by PEG-mediated transfer into protoplasts of the wild type.pZESH1,a green fluorescent expression vector containing FolSid1 gene was constructed,and the subcellular localization of FolSid1-EGFP fusion protein was performed.The results showed that the sequence of FolSid1 gene consisted of 5 392 bp,which contained 3 introns.Compared with the wild type and the external insertion mutant, the knockout mutant ΔFolSid1 conidia showed a significant decrease in yield, although they did not differ in morphology and size; morphological observations revealed that the growth rate of colonies from the knockout mutant was significantly slower. The experiments of subcellular localization showed that FolSid1 protein was located in the cell membrane of mycelia cell.FolSid1 gene regulated the vegetative growth of mycelium,conidiogenesis and pathogenicity of Fusarium oxysporum f.sp.lini.

Key words: Fusarium oxysporum f.sp.lini, FolSid1, Split Marker, Gene disruption

高芳, 侯占铭. 尖孢镰刀菌亚麻专化型FolSid1基因敲除及功能分析[J]. 华北农学报, 2025, 40(1): 189-198. doi: 10.7668/hbnxb.20194757.

GAO Fang, HOU Zhanming. Knockout and Function Analysis of FolSid1 Gene in Fusarium oxysporum f.sp.lini[J]. Acta Agriculturae Boreali-Sinica, 2025, 40(1): 189-198. doi: 10.7668/hbnxb.20194757.

http://www.hbnxb.net/CN/Y2025/V40/I1/189

图/表 12

表1 引物序列

Tab.1 Primer sequence

引物名称 Primer name	引物序列(5'—3') Primer sequence (5'—3')
FolSid1 1F	CTTGTTGCGTTGCATGATC
FolSid1 2R	CAGCTTCGTGGAGTCGTC
FolSid1 3F	CCGGTACCTTGCCTTAGTG
FolSid1 4R	CAGTTTCGCCAGTTGCC
FolSid1 5F	CGTTGCTGAGCATTATCAGG
FolSid1 6R	GAAGTCCTTGCCGAAGTTG
FolSid1 7F	GCCAACTTCAGCGAAAC
FolSid1 8R	GGAGTATCAGCATGACGG
FolSid1 9F	CGAGCTGCATGCTCAAAC
FolSid1 10R	GGAACGCCGATATGTG
FolSid1 11R	GACAAGATATCGCCGTGG
FolSid1 12R	GAAGAGAACAACGCCTACAAG
FolSid1 KO1F	GGCGTGATATGTGATGTGATATG
FolSid1 KO2R	TTGACCTCCACTAGCTCCAGCCAAGCCG GTAGTCCTGACCGTTGTCTGG
FolSid1 KO3F	ATAGAGTAGATGCCGACCGCGGGTTCC GTGCACAAGGCGTTAAAGG
FolSid1 KO4R	CGATACGCGGTACGCCGAG
HYG-F	GGCTTGGCTGGAGCTAGTGGAGGTCAA
HY-R	TTCGGACCGCAAGGAATCGGTCAATAC
YG-F	GATGTAGGAGGGCGTGGATATGTCCT
HYG-R	ATAGAGTAGATGCCGACCGCGGGTTC
FolSid1 N1F	CGTTGCTGAGCATTATCAGGTCC
FolSid1 N2R	CGCATAGTCGATCCAGAGTTGCC
FolSid1 C1F	CCAGACAACGGTCAGGACTACC
FolSid1 C2R	GACGATGGCTTCCGGTGCTGC
FolSid1 G1F	GCCTGCAGGTCGACTCTAGAGGATCCGC CAACCATCGACGATCGAACC
FolSid1 G2R	CAGCTCCTCGCCCTTGCTCACCATAACG GCAGGCCATCGAAGGCGAAGAC
Neo F	GAGAGGCTATTCGGCTATGACT
Neo R	GGCCACAGTCGATGAATCCAGA
FolSid1-F	GACCATGACTCCTAAGGACGAGC
EGFP-R	TGAAGCACTGCACGCCGT

图1 4种真菌氨基酸序列比对

Fig.1 Amino acid sequence alignment of four fungi

图2 根据10种真菌氨基酸序列构建的系统进化树

Fig.2 Phylogenetic tree based on amino acid sequences of 10 fungi

图3 FolSid1基因缺失盒的构建 M.Marker Ⅲ。A.FolSid1上、下游序列PCR扩增产物(1.FolSid1上游PCR扩增产物;2.FolSid1下游PCR扩增产物)。B.hph上、下游断裂片段PCR扩增产物(1.hph上游断裂片段PCR扩增产物;2.hph下游断裂片段PCR扩增产物)。C.FolSid1 Split Marker重叠PCR产物(1.FolSid1 1F/HY-R重叠PCR产物;2.YG-F/FolSid1 4R重叠PCR产物)。

Fig.3 Construction of gene deletion box of FolSid1 M.Marker Ⅲ.A.PCR amplification product of upstream and downstream sequence of FolSid1(1.PCR amplification product of upstream sequence of FolSid1; 2.PCR amplification product of downstream sequence of FolSid1).B.PCR amplification product of upstream and downstream fragments of hph;(1.PCR amplification product of upstream fragment of hph; 2.PCR amplification product of downstream fragment of hph).C.Overlapping PCR product for FolSid1 Split Marker;(1.FolSid1 1F/HY-R overlapping PCR product; 2.YG-F/FolSid1 4R overlapping PCR product).

图4 FolSid1转化子的筛选与鉴定 A.ΔFolSid1基因组DNA(M.1 kb plus DNA ladder,1—5.转化子1—5基因组DNA);B.以FolSid1 N1F/FolSid1 N2R为引物对FolSid1敲除突变体进行阴性筛选;C.以FolSid1 KO1F/HY-R为引物对FolSid1敲除突变体进行阳性筛选;D.以HYG-F/HYG-R为引物对FolSid1敲除突变体进行阳性筛选;E.以YG-F/FolSid1 KO4R为引物对FolSid1敲除突变体进行阳性筛选。B—E:M.Marker Ⅲ;1—4.以FolSid1转化子DNA为模板,5.以hm DNA为模板。

Fig.4 Screening and identification of FolSid1 transformants A.ΔFolSid1 genomic DNA(M.1 kb plus DNA ladder; 1—5.Transformants 1—5 genomic DNA);B.Negative screening of FolSid1 knockout mutants using FolSid1 N1F/FolSid1 N2R as primers; C.Positive screening of FolSid1 knockout mutants using FolSid1 KO1F/HY-R as primers;D.Positive screening of FolSid1 knockout mutants using HYG-F/HYG-R as primers; E.Positive screening of FolSid1 knockout mutants using YG-F/FolSid1 KO4R as primers.B-E:M.Marker Ⅲ;1-4.FolSid1 transformant DNA was used as template;5.hm DNA was used as a template.

图5 不同菌株表型分析 A.野生型hm、ΔFolSid1和ecFolSid1在V8固体培养基上培养的菌落形态;B.hm、ΔFolSid1和ecFolSid1的菌丝生长速度;C.hm、ΔFolSid1和ecFolSid1在YEPD液体培养基中的菌丝形态;D.hm、ΔFolSid1和ecFolSid1在CMC液体培养基中的分生孢子产量。

Fig.5 Phenotypic analysis of different strains A.Colony morphology of wild type hm,ΔFolSid1and ecFolSid1 cultured on V8 solid medium; B.The mycelium growth rate of hm,ΔFolSid1and ecFolSid1; C.Hyphal morphology of hm,ΔFolSid1and ecFolSid1 in YEPD fluit medium; D.Conidial production of hm,ΔFolSid1and ecFolSid1 in CMC fluit medium.

表2 不同菌株中分生孢子的产量

Tab.2 Conidial production in different strains

菌株 Strain	分生孢子量/(×10⁶/mL) Conidia quantity
hm	7.545±2.339 8a
ΔFolSid1	3.572±0.721 9b
ecFolSid1	6.628±1.060 0a

图6 不同菌株的分生孢子悬浮液对亚麻幼苗的侵染 A.不同菌株分生孢子悬浮液对亚麻幼苗侵染的正视图;B.不同菌株分生孢子悬浮液对亚麻幼苗侵染的俯视图。

Fig.6 Infection of flax seedlings by conidia suspension of different strains A.Positive view of infection of different strains conidia suspensions on flax seedlings; B.Top view of infection of different strains conidia suspensions on flax seedlings.

图7 质粒 pZESH1 的构建 A.pZEH1质粒DNA;B.线性化载体pZEH1;C.以FolSid1基因组DNA为模板、FolSid1 G1F/FolSid1 G2R为引物的PCR扩增产物。M.1 kb plus DNA ladder。

Fig.7 Construction of plasmid pZESH1 A.pZEH1 plasmid DNA; B.Linearized vector pZEH1; C.The PCR amplification product using FolSid1 genomic DNA as template and FolSid1 G1F/FolSid1 G2R as primer.M.1 kb plus DNA ladder.

图8 质粒 pZESH1 的筛选 A.以不同菌株为模板、FolSid1 N1F/FolSid1 N2R为引物的PCR扩增产物;B.菌落12、19质粒DNA;C.Xba Ⅰ 酶切质粒pZESH1;D.以质粒DNA为模板、FolSid1 N1F/FolSid1 N2R为引物的PCR扩增产物;E.以质粒DNA为模板、以FolSid1-F/EGFP-R为引物的PCR扩增产物。M.1 kb plus DNA ladder(A—C);Marker Ⅲ(D—E)。

Fig.8 Screening of plasmid pZESH1 A.The PCR amplification product using different strains as template and FolSid1 N1F/FolSid1 N2R as primer; B.Colony 12,19 plasmid DNA; C.Xba Ⅰ enzyme digestion recombinant plasmid pZESH1; D.The PCR amplification product using plasmid DNA as template and FolSid1 N1F/FolSid1 N2R as primer; E.The PCR amplification product using plasmid DNA as template and FolSid1-F/EGFP-R as primer.M.1 kb plus DNA ladder(A—C);Marker Ⅲ(D—E).

图9 亚细胞定位转化子的筛选 A.亚细胞定位转化子DNA;B.以质粒DNA为模板、Neo F/Neo R为引物的PCR扩增产物;C.以质粒DNA为模板、FolSid1-F/EGFP-R为引物的PCR扩增产物。M.1 kb plus DNA ladder(A);Marker Ⅲ(B—C)。

Fig.9 Screening of subcellular locatable transformants A.Transformant DNA of subcellular localization; B.The PCR amplification product using plasmid DNA as template and Neo F/Neo R as primer; C.The PCR amplification product using plasmid DNA as template and FolSid1-F/EGFP-R as primer.M.1 kb plus DNA ladder(A); Marker Ⅲ(B—C).

图10 FolSid1-EGFP蛋白的亚细胞定位 A.FolSid1-egfp菌株,箭头所指处为菌丝体细胞膜位置;B.hm菌株。

Fig.10 Subcellular localization of FolSid1-EGFP protein A.FolSid1-egfp strain,arrows indicate the location of mycelium cell membrane; B.hm strain.

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尖孢镰刀菌亚麻专化型FolSid1基因敲除及功能分析

Knockout and Function Analysis of FolSid1 Gene in Fusarium oxysporum f.sp.lini

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尖孢镰刀菌亚麻专化型FolSid1基因敲除及功能分析

Knockout and Function Analysis of FolSid1 Gene in Fusarium oxysporum f.sp.lini

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