思茅松毛虫核型多角体病毒及全基因组分析
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摘要
松毛虫类害虫是我国重大森林害虫之一。思茅松毛虫Dendrolimus kikuchiiMatsumura属鳞翅目Lepidoptera枯叶蛾科Lasiocampidae松毛虫属Dendrolimus。目前防治该害虫以化学农药为主。杆状病毒具有很强的专化性，对环境和非靶标生物安全，是一种理想的病原微生物杀虫剂。世界上已有50多种商品化生产的杆状病毒杀虫剂用于农林害虫的控制。因此，寻找致病力强的杆状病毒来防治该害虫显得非常必要。目前，作者在云南思茅松毛虫重灾区采集到了一株新的思茅松毛虫核型多角体病毒，主要研究结果如下：
1.思茅松毛虫核型多角体病毒（DekiNPV）形态学和毒力研究
发现了一株新的思茅松毛虫核型多角体病毒，定名为DekiNPV。多角体呈不规则型，直径大小0.79~2.31μm（n=100），平均直径1.64±0.1μm，其表面凹凸不平，且有大量的长条形、圆形孔洞，长条形居多，大小（173.00~254.00）×(55.10~116.00) nm；一个多角体中有多个杆状病毒粒子，杆状病毒粒子长约（252.22~359.38）×（70.18~200.00）nm，两端钝圆或平截，其内含有1~9个核衣壳，大小（24.00~28.60）×（242.00~340.00）nm,平均大小25.80±0.86×265.00±12.66nm (n=50)，为多粒包埋型。用浓度2.2×104~2.2×108PIB/mL5个浓度的DekiNPV接种于3龄思茅松毛虫幼虫发现，DekiNPV对3龄思茅松毛虫幼虫有很强的毒力；LT50随着浓度的降低而增长，分别为6.89、7.18、8.16、10.31和11.13d，LC50为1.72×105PIB/mL；3龄思茅松毛虫幼虫感染DekiNPV后第7d幼虫开始大量死亡，死亡高峰期为感染病毒后的第7~10d。对在实验室连续传四代后的DekiNPV进行超微结构观察发现，传代后的DekiNPV形态结构与原代一致，具有稳定性。
2. DekiNPV基因组全序列分析
分析了DekiNPV全基因组序列，DekiNPV基因组大小为141，454bp， C+G含量为48.04%。预测DekiNPV含有146个开放阅读框(ORF)，其大小在150个核苷酸以上，并与其它ORF有最小的重叠。其中有133个DekiNPV的ORF与报道已测序的杆状病毒具有同源性，13个ORF为该病毒特有，占全基因组的12.4%。基于DekiNPV的29个杆状病毒核心基因构建了该病毒的进化树，进化分析发现DekiNPV属于鳞翅目杆状病毒Group Ⅰ组，与MaviMNPV、BomaNPV、BmNPV、PlxyMNPV、RoMNPV和AcMNPV亲缘关系近。DekiNPV基因组中含有16个保守的结构基因，DekiNPV基因组缺失P6.9和odv-e56。该基因含有抗凋亡基因iap-1，iap-2和iap-3。DekiNPV基因组含中有12个bro基因。DekiNPV分别与AcMNPV、BmNPV、ChchNPV、LdMNPV、MacoNPV-B、SeMNPV和XecnGV进行了基因对等比较分析，发现该病毒与AcMNPV和BmNPV表现出了最好的线性关系。同源区分析发现，DekiNPV中共有11个hr。
3. DekiNPV的PCR检测方法
根据DekiNPV特有阅读框ORF146设计两对引物建立PCR检测方法，用DekiNPV基因组DNA为模板扩增出了426bp和655bp片段，PCR产物经测序结果与引物选取片段大小一致。分别以两对引物对DekiNPV的基因组DNA、多角体进行检测，最小量检测均达到了0.8fg/mL DNA和5PIB/mL，在感病虫体内检测到了DekiNPV的存在。该方法具有灵敏度高、特异性强、早期检测等特点，可以用于DekiNPV寄主域、替代寄主和病毒在种群中动态变化等领域的研究。
4. DekiNPV寄主域及其替代寄主的研究
用11种鳞翅目昆虫（云南松毛虫Dendrolimus houi、马尾松毛Dendrolimus punctatus、文山松毛虫Dendrolimus punctatus wenshanensis、赤松毛虫Dendrolimus spectabilis、美国白蛾Hyphantria cunea、舞毒蛾Lymantria dispar、灰茶尺蠖Ectropis grisescens、棉铃虫Helicoverpa armigera、甜菜夜蛾Spodoptera exigua、小菜蛾Plutella xylostella和斜纹夜蛾Spodoptera litura）和两株昆虫细胞系[sf-9细胞和杨扇舟蛾细胞（CAF-clan II）]研究了DekiNPV的寄主域和替代寄主。研究发现DekiNPV具有很强的专化性，但DekiNPV可以诱发美国白蛾、云南松毛虫、马尾松毛、文山松毛虫和赤松毛虫体内潜伏性感染病毒。
Dendrolimus kikuchii (Matsumura)(Lepidoptera: Lasiocampidae), is a serious forestpest for a variety of conifers. In China, D. kikuchii mainly occurs in the south part andchemical pesticides were primely used as the first control method. However, chemicalinsecticides may poison the non-target organisms (e.g., humans, livestock and natural enemy),pollute environment, and induce insecticide-resistant pests. Baculoviruses are ideal tools inintegrated pest management because they are usually highly specific to their host insects anddo not affect other arthropods, including pest predators and parasitoids. They also pose nohazard to the environment, vertebrates, plants, and natural enemies. Over50baculovirusproducts have been used to control agricultural and forestry pests worldwide. Therefore, thescreening of high toxicity and strong stability virus is an imperative work for the biologicalcontrol of D. kikuchii.
In present study, a nucleopolyhedrovirus was isolated from the infected larvae of D.kikuchii, which is a serious pest for a variety of conifers in China. The main results are asfollows:
1. Morphological characteristics and virulence of D. kikuchii nucleopolyhedrovirus(DekiNPV)
A new strain of nucleopolyhedrovirus has been isolated from infected larvae of the D.kikuchii (Matsumura)(Lepidoptera: Lasiocampidae) in China and named as DekiNPV. Thescanning electron micrograph results showed that OBs of D. kikuchii are mostly polyhedralshape,0.79~2.31μm in diameter (n=100), average diameter is (1.64±0.1) μm, with a lots ofsmall holes on their surface,(173.00~254.00)×(55.10~116.00) nm in size. Ultrathin sectionsrevealed that each OB contains many virions,(252.22~359.38)×(70.18~200.00) nm in size.The virion is rod-shaped with the truncated or obtuse ends, consists of multiple nucleocapsids(up to9) within a single viral envelope. The size of nucleocapsid is(24.00~28.60)×(242.00~340.00) nm in size, and average25.80±0.86nm (mean±SE) inwidth and265.00±12.66nm in length (n=50). The results of infection study showed thatDekiNPV could infect and kill the third-larvae of D. kikuchii. LC50was1.72×105PIB/mL. At the tested five concentrations (from2.2×104to2.2×108PIB/mL), LT50were6.89,7.18,8.16,10.31and11.13days, respectively. A large number of D. kikuchii larvae begin to die from theseventh day of infection, and peak mortality from the senventh to tenth days. Our researchdemonstrates that the virus has been propagated4generations in the fourth-instars larvae of D.kikuchii has stable ultrastructural morphological.
2. Genomic sequencing and analysis of DekiNPV
The complete genome of DekiNPV from an important pest of pine, D. kikuchiiMatsumura (Lepidoptera: Lasiocampidae), was sequenced and analysied. The DekiNPVgenome was141,454bp in size with C+G content of48.04%. It is predicted to contain146open reading frames (ORFs).133DekiNPV ORFs are homologues of previously reportedbaculovirus genes, and13ORFs are unique to DekiNPV, accounting for12.4%of the genomein total. A phylogenetic tree was constructed based on29core baculovirus genes. TheDekiNPV grouped into the lepidopteran-specific NPV (Group Ⅰ), and had a closerelationship to MaviMNPV, BomaNPV, BmNPV, PlxyMNPV, RoMNPV and AcMNPV. Thisanalysis also suggested that the DekiNPV is more primitive than those6NPVs of the sameclade. The DekiNPV genome contains16conserved structural genes，and absent P6.9andodv-e56. The DekiNPV genome contains apoptosis inhibiting gene, such as iap-1，iap-2andiap-3. The result of the identity-GeneParity analysis of SeMNPV and DekiNPV, LdMNPVand DekiNPV, MacoNPV-B and DekiNPV, ChchNPV and DekiNPV, AcMNPV andDekiNPV, BmNPV and DekiNPV, and XecnGV and DekiNPV, which compares all both theoverall gene arrangement and the homology between genes. It was show that AcMNPV andBmNPV appear to be the most collinear with DekiNPV, but they most genes are inverted withthe genes of DekiNPV. There are11hrs in DekiNPV genome.
3. Polymerase chain reaction assay for the detection of D. kikuchii nucleopolyhedrovirus
A polymerase chain reaction (PCR) assay was developed with a pair of primers, designedbased on the unique open reading frame (ORF)146of DekiNPV. DekiNPV DNA served astemplate to amplify426bp and655bp segments. Results from the sequenced PCR productswere consistent with the size of selected primer segments. Two primers were applied to assayDekiNPV DNA and detect intact DekiNPV occlusion bodies (OBs), showing a minimumdetection of0.8fg/mL DNA and5OBs/mL, respectively, indicating the existence ofDekiNPV in the infected insects. The assay is characterized with high sensitivity, remarkablespecificity and early assay, demonstrating its potential for applications in the studies ofDekiNPV host range, substitution of DekiNPV hosts, dynamic DekiNPV changes in D.kikuchii population and routine molecular diagnostic procedures.4. Host range and substitution of DekiNPV
We examined the infectivity and replication of DekiNPV in11other lepidopteran species,including Dendrolimus houi, Dendrolimus punctatus, Dendrolimus punctatus wenshanensis,Dendrolimus spectabilis, Hyphantria cunea, Lymantria dispar, Ectropis grisescens,Helicoverpa armigera, Spodoptera exigua, Plutella xylostella, Spodoptera litura larvae, sf-9cell, and Clostera anachoreta cell (CAF-clan II) to define the host range of DekiNPV andidentify its suitable alternate hosts. Our study showed that DekiNPV is highly specific to D.kikuchii, but triggers covert infections in H. cunea, D. houi, D. punctatus, D. punctatuswenshanensis, and D. spectabilis larvae into overt symptoms, indicating that DekiNPV issuitable as an ideal biocontrol agent for D. kikuchii, D. houi, D. punctatus, D. punctatuswenshanensis, D. spectabilis, and H. cunea.
1.思茅松毛虫核型多角体病毒（DekiNPV）形态学和毒力研究
发现了一株新的思茅松毛虫核型多角体病毒，定名为DekiNPV。多角体呈不规则型，直径大小0.79~2.31μm（n=100），平均直径1.64±0.1μm，其表面凹凸不平，且有大量的长条形、圆形孔洞，长条形居多，大小（173.00~254.00）×(55.10~116.00) nm；一个多角体中有多个杆状病毒粒子，杆状病毒粒子长约（252.22~359.38）×（70.18~200.00）nm，两端钝圆或平截，其内含有1~9个核衣壳，大小（24.00~28.60）×（242.00~340.00）nm,平均大小25.80±0.86×265.00±12.66nm (n=50)，为多粒包埋型。用浓度2.2×104~2.2×108PIB/mL5个浓度的DekiNPV接种于3龄思茅松毛虫幼虫发现，DekiNPV对3龄思茅松毛虫幼虫有很强的毒力；LT50随着浓度的降低而增长，分别为6.89、7.18、8.16、10.31和11.13d，LC50为1.72×105PIB/mL；3龄思茅松毛虫幼虫感染DekiNPV后第7d幼虫开始大量死亡，死亡高峰期为感染病毒后的第7~10d。对在实验室连续传四代后的DekiNPV进行超微结构观察发现，传代后的DekiNPV形态结构与原代一致，具有稳定性。
2. DekiNPV基因组全序列分析
分析了DekiNPV全基因组序列，DekiNPV基因组大小为141，454bp， C+G含量为48.04%。预测DekiNPV含有146个开放阅读框(ORF)，其大小在150个核苷酸以上，并与其它ORF有最小的重叠。其中有133个DekiNPV的ORF与报道已测序的杆状病毒具有同源性，13个ORF为该病毒特有，占全基因组的12.4%。基于DekiNPV的29个杆状病毒核心基因构建了该病毒的进化树，进化分析发现DekiNPV属于鳞翅目杆状病毒Group Ⅰ组，与MaviMNPV、BomaNPV、BmNPV、PlxyMNPV、RoMNPV和AcMNPV亲缘关系近。DekiNPV基因组中含有16个保守的结构基因，DekiNPV基因组缺失P6.9和odv-e56。该基因含有抗凋亡基因iap-1，iap-2和iap-3。DekiNPV基因组含中有12个bro基因。DekiNPV分别与AcMNPV、BmNPV、ChchNPV、LdMNPV、MacoNPV-B、SeMNPV和XecnGV进行了基因对等比较分析，发现该病毒与AcMNPV和BmNPV表现出了最好的线性关系。同源区分析发现，DekiNPV中共有11个hr。
3. DekiNPV的PCR检测方法
根据DekiNPV特有阅读框ORF146设计两对引物建立PCR检测方法，用DekiNPV基因组DNA为模板扩增出了426bp和655bp片段，PCR产物经测序结果与引物选取片段大小一致。分别以两对引物对DekiNPV的基因组DNA、多角体进行检测，最小量检测均达到了0.8fg/mL DNA和5PIB/mL，在感病虫体内检测到了DekiNPV的存在。该方法具有灵敏度高、特异性强、早期检测等特点，可以用于DekiNPV寄主域、替代寄主和病毒在种群中动态变化等领域的研究。
4. DekiNPV寄主域及其替代寄主的研究
用11种鳞翅目昆虫（云南松毛虫Dendrolimus houi、马尾松毛Dendrolimus punctatus、文山松毛虫Dendrolimus punctatus wenshanensis、赤松毛虫Dendrolimus spectabilis、美国白蛾Hyphantria cunea、舞毒蛾Lymantria dispar、灰茶尺蠖Ectropis grisescens、棉铃虫Helicoverpa armigera、甜菜夜蛾Spodoptera exigua、小菜蛾Plutella xylostella和斜纹夜蛾Spodoptera litura）和两株昆虫细胞系[sf-9细胞和杨扇舟蛾细胞（CAF-clan II）]研究了DekiNPV的寄主域和替代寄主。研究发现DekiNPV具有很强的专化性，但DekiNPV可以诱发美国白蛾、云南松毛虫、马尾松毛、文山松毛虫和赤松毛虫体内潜伏性感染病毒。
Dendrolimus kikuchii (Matsumura)(Lepidoptera: Lasiocampidae), is a serious forestpest for a variety of conifers. In China, D. kikuchii mainly occurs in the south part andchemical pesticides were primely used as the first control method. However, chemicalinsecticides may poison the non-target organisms (e.g., humans, livestock and natural enemy),pollute environment, and induce insecticide-resistant pests. Baculoviruses are ideal tools inintegrated pest management because they are usually highly specific to their host insects anddo not affect other arthropods, including pest predators and parasitoids. They also pose nohazard to the environment, vertebrates, plants, and natural enemies. Over50baculovirusproducts have been used to control agricultural and forestry pests worldwide. Therefore, thescreening of high toxicity and strong stability virus is an imperative work for the biologicalcontrol of D. kikuchii.
In present study, a nucleopolyhedrovirus was isolated from the infected larvae of D.kikuchii, which is a serious pest for a variety of conifers in China. The main results are asfollows:
1. Morphological characteristics and virulence of D. kikuchii nucleopolyhedrovirus(DekiNPV)
A new strain of nucleopolyhedrovirus has been isolated from infected larvae of the D.kikuchii (Matsumura)(Lepidoptera: Lasiocampidae) in China and named as DekiNPV. Thescanning electron micrograph results showed that OBs of D. kikuchii are mostly polyhedralshape,0.79~2.31μm in diameter (n=100), average diameter is (1.64±0.1) μm, with a lots ofsmall holes on their surface,(173.00~254.00)×(55.10~116.00) nm in size. Ultrathin sectionsrevealed that each OB contains many virions,(252.22~359.38)×(70.18~200.00) nm in size.The virion is rod-shaped with the truncated or obtuse ends, consists of multiple nucleocapsids(up to9) within a single viral envelope. The size of nucleocapsid is(24.00~28.60)×(242.00~340.00) nm in size, and average25.80±0.86nm (mean±SE) inwidth and265.00±12.66nm in length (n=50). The results of infection study showed thatDekiNPV could infect and kill the third-larvae of D. kikuchii. LC50was1.72×105PIB/mL. At the tested five concentrations (from2.2×104to2.2×108PIB/mL), LT50were6.89,7.18,8.16,10.31and11.13days, respectively. A large number of D. kikuchii larvae begin to die from theseventh day of infection, and peak mortality from the senventh to tenth days. Our researchdemonstrates that the virus has been propagated4generations in the fourth-instars larvae of D.kikuchii has stable ultrastructural morphological.
2. Genomic sequencing and analysis of DekiNPV
The complete genome of DekiNPV from an important pest of pine, D. kikuchiiMatsumura (Lepidoptera: Lasiocampidae), was sequenced and analysied. The DekiNPVgenome was141,454bp in size with C+G content of48.04%. It is predicted to contain146open reading frames (ORFs).133DekiNPV ORFs are homologues of previously reportedbaculovirus genes, and13ORFs are unique to DekiNPV, accounting for12.4%of the genomein total. A phylogenetic tree was constructed based on29core baculovirus genes. TheDekiNPV grouped into the lepidopteran-specific NPV (Group Ⅰ), and had a closerelationship to MaviMNPV, BomaNPV, BmNPV, PlxyMNPV, RoMNPV and AcMNPV. Thisanalysis also suggested that the DekiNPV is more primitive than those6NPVs of the sameclade. The DekiNPV genome contains16conserved structural genes，and absent P6.9andodv-e56. The DekiNPV genome contains apoptosis inhibiting gene, such as iap-1，iap-2andiap-3. The result of the identity-GeneParity analysis of SeMNPV and DekiNPV, LdMNPVand DekiNPV, MacoNPV-B and DekiNPV, ChchNPV and DekiNPV, AcMNPV andDekiNPV, BmNPV and DekiNPV, and XecnGV and DekiNPV, which compares all both theoverall gene arrangement and the homology between genes. It was show that AcMNPV andBmNPV appear to be the most collinear with DekiNPV, but they most genes are inverted withthe genes of DekiNPV. There are11hrs in DekiNPV genome.
3. Polymerase chain reaction assay for the detection of D. kikuchii nucleopolyhedrovirus
A polymerase chain reaction (PCR) assay was developed with a pair of primers, designedbased on the unique open reading frame (ORF)146of DekiNPV. DekiNPV DNA served astemplate to amplify426bp and655bp segments. Results from the sequenced PCR productswere consistent with the size of selected primer segments. Two primers were applied to assayDekiNPV DNA and detect intact DekiNPV occlusion bodies (OBs), showing a minimumdetection of0.8fg/mL DNA and5OBs/mL, respectively, indicating the existence ofDekiNPV in the infected insects. The assay is characterized with high sensitivity, remarkablespecificity and early assay, demonstrating its potential for applications in the studies ofDekiNPV host range, substitution of DekiNPV hosts, dynamic DekiNPV changes in D.kikuchii population and routine molecular diagnostic procedures.4. Host range and substitution of DekiNPV
We examined the infectivity and replication of DekiNPV in11other lepidopteran species,including Dendrolimus houi, Dendrolimus punctatus, Dendrolimus punctatus wenshanensis,Dendrolimus spectabilis, Hyphantria cunea, Lymantria dispar, Ectropis grisescens,Helicoverpa armigera, Spodoptera exigua, Plutella xylostella, Spodoptera litura larvae, sf-9cell, and Clostera anachoreta cell (CAF-clan II) to define the host range of DekiNPV andidentify its suitable alternate hosts. Our study showed that DekiNPV is highly specific to D.kikuchii, but triggers covert infections in H. cunea, D. houi, D. punctatus, D. punctatuswenshanensis, and D. spectabilis larvae into overt symptoms, indicating that DekiNPV issuitable as an ideal biocontrol agent for D. kikuchii, D. houi, D. punctatus, D. punctatuswenshanensis, D. spectabilis, and H. cunea.
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