抗虫水稻对褐飞虱抗性研究及褐飞虱apterousA基因的功能研究
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摘要
褐飞虱是亚洲许多国家水稻上的重要害虫,在我国长江流域及华南流域和西南广大稻区都有分布,能远距离迁飞,繁殖力惊人,常暴发成灾。为揭示褐飞虱灾变的内在机制,必须借助于现代分子生物学手段探索其生长发育的生命特征,开展相关基因的功能研究。本文旨在通过抗虫水稻对褐飞虱生长发育的影响,探索抗虫水稻对褐飞虱的抗虫机制；同时克隆褐飞虱生长发育相关基因apterousA,分析其功能,为褐飞虱的防控提供了新的靶标基因。
一、含Bph14或Bph15基因抗虫水稻对褐飞虱抗性的研究
1.苗期抗虫鉴定及分蘖期蜜露量测定结果
MH63::14, MH63::15和MH63：：14&15水稻品系对褐飞虱的抗性级别均小于3,对褐飞虱表现为高抗,并且取食3种抗虫水稻的褐飞虱所分泌的蜜露量显著低于非抗性水稻上褐飞虱的蜜露量。结果表明,含抗虫基因的水稻在苗期和分蘖期对褐飞虱均有较强抗性。
2.抗虫水稻对褐飞虱生长发育的影响
与MH63相比,MH63：：15和MH63：：14&15水稻品系延长了褐飞虱若虫的发育历期,缩短了褐飞虱雌成虫的寿命,降低了褐飞虱的短翅率、雌虫比率、交配率和产卵量。结果表明,抗虫水稻不利于褐飞虱的生长发育及繁殖。
3.抗虫水稻对褐飞虱抗生性的影响
抗虫水稻MH63：：15和MH63：：14&15上累计褐飞虱若虫数和雌成虫数均显著低于非抗虫品种MH63上相应褐飞虱数量,并且褐飞虱雌成虫在抗虫分蘖期水稻上的产卵量也显著低于对照,表明抗虫水稻对褐飞虱有很好的抗生性。
4.抗虫水稻对褐飞虱体内P450基因的表达调控
本章节研究了取食抗虫水稻后褐飞虱体内部分P450基因的表达情况,发现取食过MH63：：14&15和MH63：：15抗性水稻后的褐飞虱体内CYP4C61, CYP4C62和CYP6CW1的表达量升高,CYP6CS1的表达量降低,而CYP303A1的表达量不变。结果表明,CYP4C61,CYP4C62和CYP6CW1在抗虫水稻抵抗褐飞虱的过程中发挥重要作用。
二、褐飞虱apterouA基因的克隆及其在翅和生殖发育中的功能研究
1.褐飞虱apterousA基因的克隆与分析
采用PCR、RT-PCR、RACE等方法克隆了褐飞虱apterousA (NlapA)基因,并根据其cDNA序列推导出氨基酸序列,GenBank的登录号为KC978728。克隆到的褐飞虱NlapA基因cDNA序列长为3004bp,开放阅读框为58-1326bp,共编码422个氨基酸,推测蛋白的分子量为46.6kDa。ApterousA蛋白有2个LIM结构域和1个homeodomain保守结构域。
2. NlapA在褐飞虱中的时空表达
研究发现,NlapA基因在褐飞虱各发育时期均有表达,若虫1-5龄和成虫1天的表达量高于成虫2-8天的表达量,并且NlapA基因在褐飞虱3龄若虫中的表达量最高。此外,NlapA基因在褐飞虱若虫的各组织中也均有表达,但在翅芽中的表达量最高,是其它组织中NlapA基因表达量的2.2-9倍。
3. NlapA RNAi后褐飞虱体内靶基因表达量下降
根据NlapA基因序列设计3段不同dsapA,将dsapA直接注射到褐飞虱的血腔里,各处理存活率之间无明显差异。取NlapA RNAi后不同时期的褐飞虱和卵,检测NlapA的表达量,结果发现,随着NlapA RNAi时间的延长,靶基因的表达量逐渐降低,第10天时,其表达量最低,仅为对照的9%;第18天时,处理组褐飞虱卵中NlapA的表达量仅为对照的19%。
4. NlapA RNAi导致褐飞虱翅畸形
NlapA RNAi后褐飞虱翅出现畸形,其翅畸形率在93%以上,而对照GFP RNAi后的褐飞虱无畸形翅产生。翅畸形主要表现在以下3个方面：表型Ⅰ：褐飞虱的翅脉弯曲导致双翅不能自然合拢；表型Ⅱ：褐飞虱翅脉上的刚毛消失,翅脉变淡；表型Ⅲ:褐飞虱翅脉之间的膜质区域变薄。5. NlapA RNAi降低褐飞虱生殖力
NlapA RNAi显著降低褐飞虱的产卵量,仅为对照产卵量的15.67-23.13%;NlapA RNAi后褐飞虱卵的孵化率为0,而对照卵孵化率为92.7%。尽管NlapA阻滞了褐飞虱卵的发育,但未影响褐飞虱卵巢的形态发育。
综上所述,含Bph14或Bph15基因抗虫水稻不利于褐飞虱的生长发育,削弱褐飞虱生殖力；NlapA RNAi导致褐飞虱翅畸形出现,降低褐飞虱的产卵量,阻滞褐飞虱卵的发育。本研究为抗虫水稻的抗虫机制、褐飞虱翅和生殖发育基因的功能研究奠定了坚实的理论基础。
The brown planthopper (BPH), Nilaparvata lugens (Stal) are serious pests in rice fields in many areas of Asia, they are distributed mainly over the Yangtze River, South China and southwestern China. The reproduction of BPH is amazing, they also can migrate to other areas and cause further damage. To illustrate inherent mechanism of disasters caused by BPH, it is necessary to study the molecular mechanism of growth and development of BPH, by molecular biological technology. In this study, the biological effects of insect-resistant rice on BPH was investigated, and that was about to expound insect-resistant mechanism of insect-resistant varieties. Fuhthermore, we cloned N. lugens apterousA(NlapA) gene relating to growth and development, analyzed its function, and providing new genetic resources for BPH controlling.
1. Biological effects of rice harbouring Bphl4and Bphl5on BPH
①The results of seedling resistance to BPH and honeydew measurement during tillering stage
The severity scores of MH63::14, MH63::15and MH63::14&15were<3.0, which indicated that insect-resistant rice lines have a high resistance to BPH. In addition, honeydew production of BPH on those insect-resistant lines was significantly lower than that on TN1and MH63. This indicated that the resistance level of introgression lines was obviously improved during seedling and tillering stage.
②The biological effects of insect-resistant lines on BPH
Compared with MH63, the development duration of BPH nymph was significantly retarded, the development duration of female was significantly decreased, and the brachyptery ratio, female ratio, copulation rate and fecundity were significantly decreased by MH63::15and MH63::14&15. This indicated that resistant rice lines were not beneficial to the growth and development of BPH.
③The antixenosis of resistant rice lines on BPH
The average number of nymphs and female adults settled on resistant rice lines (MH63::15and MH63::14&15) was significantly lower than that on MH63, and the number of BPH eggs per tiller was significantly lower than MH63. This indicated that the antixenosis of resistant rice lines on BPH was high.
④The expression of P450gene was regulated by insect-resistant rice in BPH
Among the five P450genes of BPH, the expression of three genes (CYP4C61, CYP4C62and CYP6CW1) was up-regulated, the expression of one gene (CYP6CS1) was down-regulated and one gene (CYP303A1) was unchanged by resistant hosts. This indicated that CYP4C61, CYP4C62and CYP6CW1were important during the insect-resistant course of resistant rice lines.
2. Gene clone and the role of NlapA in the wing development and reproduction of BPH
①Cloning and sequence analysis of NlapA
The NlapA cDNA is3004bp long and contains a unique open reading frame (ORF) spanning nucleotides58-1326bp. The polypeptide deduced from the ORF comprises422amino acides, with a calculated molecular mass of46.6kDa. The predicted protein contains two conseved LIM domains and one homeodomain.
②The expression of NlapA in different tissues and different periods during BPH development
The results show that the relative expressions of NlapA was investigated in all stages of BPH, the relative expressions of NlapA during first-fifth instar nymphs and one day after BPH emergence were higher than those during two days-eight days after BPH emergence. And the relative expression of NlapA in the third instar nymphs is the highest. In addition, the relative expressions of NlapA are stable during different tissues of BPH, but the relative expression of NlapA is the highest in the wing buds of BPH. And the relative expression of NlapA in wing buds after NlapA RNAi is2.5-6times of that in other tissues.
③The expression of NlapA in BPH after NlapA RNAi
In this study, we design three different dsapA according the sequence of NlapA, he results of NlapA RNAi shows that the relative expressions of NlapA in different periods during BPH development were significantly lower than those in control, the relative expression of NlapA is gradually decreasing over days after injection of dsapA, and the relative expression of NlapA in BPH is the lowest on the tenth day after injection of dsapA. Furthermore, the relative expression of NlapA in eggs after NlapA RNAi is significantly lower than control, and it is only19%of that in control.
④NlapA RNAi result in wing abnormality
The abnormality rates of BPH after injection of dsapAs are above93%, however, the abnormality rates of BPH after injection of dsGFP is zero. NlapA RNAi results in wing morphological defects in BPH, the main wing defects as follows:I:The veins on the wing is crooked, and that results in BPH not to form normal folded wing; Ⅱ:No sensory bristles are existed on the forewing veins, and veins become thinner; Ⅲ:The membranous regions between veins become thinner.
⑤NlapA RNAi decreased the reproduction of BPH
The fecundity of BPH after NlapA RNAi is significantly decreased by dsapA than that in control, and it is only15.67-23.13%of that in control. The emergence rate of BPH after NlapA RNAi is zero, but the emergence rate of BPH after GFP RNAi is92.7%. Although dsapA stops the development of eggs, it has no effect on the morphological characteristics of ovary.
Taken together, the results indicated that resistant rice lines were not beneficial to the growth and development of BPH, and they slao decreased the reproduction of BPH; knockdown of NlapA expression in vivo RNAi generated phenotypic defects in the wing, decreased the number of offspring and stopped the development of eggs. This technique will provide powerful tools for functional genomics of BPH, so as to lay a solid foundation of theoretical knowledge for the study of BPH development genes.
一、含Bph14或Bph15基因抗虫水稻对褐飞虱抗性的研究
1.苗期抗虫鉴定及分蘖期蜜露量测定结果
MH63::14, MH63::15和MH63：：14&15水稻品系对褐飞虱的抗性级别均小于3,对褐飞虱表现为高抗,并且取食3种抗虫水稻的褐飞虱所分泌的蜜露量显著低于非抗性水稻上褐飞虱的蜜露量。结果表明,含抗虫基因的水稻在苗期和分蘖期对褐飞虱均有较强抗性。
2.抗虫水稻对褐飞虱生长发育的影响
与MH63相比,MH63：：15和MH63：：14&15水稻品系延长了褐飞虱若虫的发育历期,缩短了褐飞虱雌成虫的寿命,降低了褐飞虱的短翅率、雌虫比率、交配率和产卵量。结果表明,抗虫水稻不利于褐飞虱的生长发育及繁殖。
3.抗虫水稻对褐飞虱抗生性的影响
抗虫水稻MH63：：15和MH63：：14&15上累计褐飞虱若虫数和雌成虫数均显著低于非抗虫品种MH63上相应褐飞虱数量,并且褐飞虱雌成虫在抗虫分蘖期水稻上的产卵量也显著低于对照,表明抗虫水稻对褐飞虱有很好的抗生性。
4.抗虫水稻对褐飞虱体内P450基因的表达调控
本章节研究了取食抗虫水稻后褐飞虱体内部分P450基因的表达情况,发现取食过MH63：：14&15和MH63：：15抗性水稻后的褐飞虱体内CYP4C61, CYP4C62和CYP6CW1的表达量升高,CYP6CS1的表达量降低,而CYP303A1的表达量不变。结果表明,CYP4C61,CYP4C62和CYP6CW1在抗虫水稻抵抗褐飞虱的过程中发挥重要作用。
二、褐飞虱apterouA基因的克隆及其在翅和生殖发育中的功能研究
1.褐飞虱apterousA基因的克隆与分析
采用PCR、RT-PCR、RACE等方法克隆了褐飞虱apterousA (NlapA)基因,并根据其cDNA序列推导出氨基酸序列,GenBank的登录号为KC978728。克隆到的褐飞虱NlapA基因cDNA序列长为3004bp,开放阅读框为58-1326bp,共编码422个氨基酸,推测蛋白的分子量为46.6kDa。ApterousA蛋白有2个LIM结构域和1个homeodomain保守结构域。
2. NlapA在褐飞虱中的时空表达
研究发现,NlapA基因在褐飞虱各发育时期均有表达,若虫1-5龄和成虫1天的表达量高于成虫2-8天的表达量,并且NlapA基因在褐飞虱3龄若虫中的表达量最高。此外,NlapA基因在褐飞虱若虫的各组织中也均有表达,但在翅芽中的表达量最高,是其它组织中NlapA基因表达量的2.2-9倍。
3. NlapA RNAi后褐飞虱体内靶基因表达量下降
根据NlapA基因序列设计3段不同dsapA,将dsapA直接注射到褐飞虱的血腔里,各处理存活率之间无明显差异。取NlapA RNAi后不同时期的褐飞虱和卵,检测NlapA的表达量,结果发现,随着NlapA RNAi时间的延长,靶基因的表达量逐渐降低,第10天时,其表达量最低,仅为对照的9%;第18天时,处理组褐飞虱卵中NlapA的表达量仅为对照的19%。
4. NlapA RNAi导致褐飞虱翅畸形
NlapA RNAi后褐飞虱翅出现畸形,其翅畸形率在93%以上,而对照GFP RNAi后的褐飞虱无畸形翅产生。翅畸形主要表现在以下3个方面：表型Ⅰ：褐飞虱的翅脉弯曲导致双翅不能自然合拢；表型Ⅱ：褐飞虱翅脉上的刚毛消失,翅脉变淡；表型Ⅲ:褐飞虱翅脉之间的膜质区域变薄。5. NlapA RNAi降低褐飞虱生殖力
NlapA RNAi显著降低褐飞虱的产卵量,仅为对照产卵量的15.67-23.13%;NlapA RNAi后褐飞虱卵的孵化率为0,而对照卵孵化率为92.7%。尽管NlapA阻滞了褐飞虱卵的发育,但未影响褐飞虱卵巢的形态发育。
综上所述,含Bph14或Bph15基因抗虫水稻不利于褐飞虱的生长发育,削弱褐飞虱生殖力；NlapA RNAi导致褐飞虱翅畸形出现,降低褐飞虱的产卵量,阻滞褐飞虱卵的发育。本研究为抗虫水稻的抗虫机制、褐飞虱翅和生殖发育基因的功能研究奠定了坚实的理论基础。
The brown planthopper (BPH), Nilaparvata lugens (Stal) are serious pests in rice fields in many areas of Asia, they are distributed mainly over the Yangtze River, South China and southwestern China. The reproduction of BPH is amazing, they also can migrate to other areas and cause further damage. To illustrate inherent mechanism of disasters caused by BPH, it is necessary to study the molecular mechanism of growth and development of BPH, by molecular biological technology. In this study, the biological effects of insect-resistant rice on BPH was investigated, and that was about to expound insect-resistant mechanism of insect-resistant varieties. Fuhthermore, we cloned N. lugens apterousA(NlapA) gene relating to growth and development, analyzed its function, and providing new genetic resources for BPH controlling.
1. Biological effects of rice harbouring Bphl4and Bphl5on BPH
①The results of seedling resistance to BPH and honeydew measurement during tillering stage
The severity scores of MH63::14, MH63::15and MH63::14&15were<3.0, which indicated that insect-resistant rice lines have a high resistance to BPH. In addition, honeydew production of BPH on those insect-resistant lines was significantly lower than that on TN1and MH63. This indicated that the resistance level of introgression lines was obviously improved during seedling and tillering stage.
②The biological effects of insect-resistant lines on BPH
Compared with MH63, the development duration of BPH nymph was significantly retarded, the development duration of female was significantly decreased, and the brachyptery ratio, female ratio, copulation rate and fecundity were significantly decreased by MH63::15and MH63::14&15. This indicated that resistant rice lines were not beneficial to the growth and development of BPH.
③The antixenosis of resistant rice lines on BPH
The average number of nymphs and female adults settled on resistant rice lines (MH63::15and MH63::14&15) was significantly lower than that on MH63, and the number of BPH eggs per tiller was significantly lower than MH63. This indicated that the antixenosis of resistant rice lines on BPH was high.
④The expression of P450gene was regulated by insect-resistant rice in BPH
Among the five P450genes of BPH, the expression of three genes (CYP4C61, CYP4C62and CYP6CW1) was up-regulated, the expression of one gene (CYP6CS1) was down-regulated and one gene (CYP303A1) was unchanged by resistant hosts. This indicated that CYP4C61, CYP4C62and CYP6CW1were important during the insect-resistant course of resistant rice lines.
2. Gene clone and the role of NlapA in the wing development and reproduction of BPH
①Cloning and sequence analysis of NlapA
The NlapA cDNA is3004bp long and contains a unique open reading frame (ORF) spanning nucleotides58-1326bp. The polypeptide deduced from the ORF comprises422amino acides, with a calculated molecular mass of46.6kDa. The predicted protein contains two conseved LIM domains and one homeodomain.
②The expression of NlapA in different tissues and different periods during BPH development
The results show that the relative expressions of NlapA was investigated in all stages of BPH, the relative expressions of NlapA during first-fifth instar nymphs and one day after BPH emergence were higher than those during two days-eight days after BPH emergence. And the relative expression of NlapA in the third instar nymphs is the highest. In addition, the relative expressions of NlapA are stable during different tissues of BPH, but the relative expression of NlapA is the highest in the wing buds of BPH. And the relative expression of NlapA in wing buds after NlapA RNAi is2.5-6times of that in other tissues.
③The expression of NlapA in BPH after NlapA RNAi
In this study, we design three different dsapA according the sequence of NlapA, he results of NlapA RNAi shows that the relative expressions of NlapA in different periods during BPH development were significantly lower than those in control, the relative expression of NlapA is gradually decreasing over days after injection of dsapA, and the relative expression of NlapA in BPH is the lowest on the tenth day after injection of dsapA. Furthermore, the relative expression of NlapA in eggs after NlapA RNAi is significantly lower than control, and it is only19%of that in control.
④NlapA RNAi result in wing abnormality
The abnormality rates of BPH after injection of dsapAs are above93%, however, the abnormality rates of BPH after injection of dsGFP is zero. NlapA RNAi results in wing morphological defects in BPH, the main wing defects as follows:I:The veins on the wing is crooked, and that results in BPH not to form normal folded wing; Ⅱ:No sensory bristles are existed on the forewing veins, and veins become thinner; Ⅲ:The membranous regions between veins become thinner.
⑤NlapA RNAi decreased the reproduction of BPH
The fecundity of BPH after NlapA RNAi is significantly decreased by dsapA than that in control, and it is only15.67-23.13%of that in control. The emergence rate of BPH after NlapA RNAi is zero, but the emergence rate of BPH after GFP RNAi is92.7%. Although dsapA stops the development of eggs, it has no effect on the morphological characteristics of ovary.
Taken together, the results indicated that resistant rice lines were not beneficial to the growth and development of BPH, and they slao decreased the reproduction of BPH; knockdown of NlapA expression in vivo RNAi generated phenotypic defects in the wing, decreased the number of offspring and stopped the development of eggs. This technique will provide powerful tools for functional genomics of BPH, so as to lay a solid foundation of theoretical knowledge for the study of BPH development genes.
引文
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