云斑天牛交配行为研究及化学感受蛋白基因的克隆
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摘要
云斑天牛Batocera horsfieldi (Hope),属于鞘翅目Coleoptera,天牛科Cerambycidae,是用材林、防护林和经济林的一种重要的毁灭性蛀干害虫。由于其幼虫生活习性隐蔽,防治十分困难。随着化学生态学的发展,基于有取食和交配等裸露活动的成虫期防治策略已经成为探索云斑天牛防治的新途径。但目前关于云斑天牛的交配行为及其交配行为所利用的信号还未见报道。为此,本研究在对云斑天牛交配行为观察的基础上,分析了嗅觉及视觉在其交配过程中的功能,并克隆了其化学感受蛋白基因,以期为阐明云斑天牛的交配行为机制及开发新的防治方法提供理论基础和技术支持。主要研究结果如下：
1.云斑天牛的交配行为
在室内对60对云斑天牛交配行为的观察发现,雄虫有80%是在没有触角接触的情况下与雌虫完成相遇进而抱对的,即雄虫主要不通过触角接触这种方式寻找雌虫；云斑天牛一次完整的交配行为分为三个阶段：相遇抱对、尝试交配和输精、配后保护。从抱对到雄虫输精之前的阶段,统称为尝试交配阶段,该阶段平均需时32.7min,在这期间雄虫有不断舔触雌虫鞘翅的行为；从雄虫开始输精到输精完成平均时间61.2s；88.3%的云斑天牛输精完成后仍然保持着抱对状态即为配后保护阶段,平均用时25.5min。
在室内对云斑天牛整个成虫期交配和产卵行为进行了观察。结果表明平均每对云斑天牛交配26.2次；交配次数第一次高峰出现在羽化出孔后的7d-8d,总的趋势是随着日龄增加交配次数减少；在一天中,交配主要发生在12：00-22：00,特别是16：00-18：00。雌虫从羽化出孔12日后产卵,产卵过程大致持续30min,平均每雌产卵39.8粒,产卵数随着日龄增加而增加,在36日龄产卵量达到峰值,之后减少进入生殖末期。
2.云斑天牛配偶定位和识别
在室内利用“Y”型嗅觉仪测试了云斑天牛雌、雄成虫之间的嗅觉反应。结果表明,以空气为对照时,雌虫对雄虫有明显的吸引作用(P<0.05),而雄虫对雌虫则无明显的吸引作用(P>0.05)；当以补充营养植物野蔷薇为对照时,雄虫对雌虫+野蔷薇、雌虫对雄虫+野蔷薇均无明显的行为趋向(P>0.05)。说明雌虫可能存在非接触性信息素,而雄虫则可能不存在非接触性信息素。
冻死雌虫与活雌虫同样能够引起测试雄虫曲腹的交配反应,但经正己烷浸泡后的雌虫则不会引起雄虫曲腹行为；而将雌虫正己烷浸泡液回滴到正己烷浸泡后的雌虫或雄虫、以及玻璃管上时,均能够引起雄虫的曲腹行为。说明云斑天牛雌虫存在接触性信息素,雄虫通过接触性信息素对雌虫进行识别。
利用气相色谱—质谱联用技术(GC-MS)对云斑天牛雌、雄成虫正己烷提取物进行了分离鉴定。在本试验条件下,云斑天牛雌、雄成虫正己烷提取物共分离鉴定出32种化合物,雌虫28种、雄虫24种；其中有8种化合物是雌虫所特有的,而对于哪种或哪些化合物对雄虫具有行为活性还需要进一步研究。
3.云斑天牛化学感受蛋白基因的克隆和序列分析
云斑天牛雄虫下颚须的舔翅行为在配偶识别方面有重要作用。为了明确其化学感受机制,利用RT-PCR技术克隆获得了云斑天牛下颚须中的化学感受蛋白基因(BhorCSP2),其核苷酸序列可读框全长384bp,编码127个氨基酸残基；推导的氨基酸序列分析表明其有保守的4个半胱氨酸位点和19个氨基酸的信号肽序列,具有昆虫化学感受蛋白的典型特征；通过对BhorCSP2编码的氨基酸和其它昆虫的30种CSP编码的氨基酸进行进化分析,发现该基因与赤拟谷盗TcasCSP11的同源性最高。
Batocera horsfieldi (Hope) (Coleoptera:Cerambycidae) is as a stem borer and a major destructive pest damaging various trees, which causing serious economic and ecology loss in China. It is difficult to control wood-boring pests because of the larvae cryptic feeding habit. With the development of chemecology, control strategies that are based on controlling or killing longhorned beetle adults which are exposed to wild for feeding and copulating. Although some species of cerambycid beetles have been studied, the mechanisms that are involved in behavior and chemical ecology of B. horsfieldi (which belongs to the lamiine) remain unknow. Research on the mating behavior and recognition mechanisms in this species will help in developing integrated pest management tools as well as increasing knowledge of this beetle. Moreover, a chemosensory protein (CSP) in maxillary palpi has also been researched to realize more about the chemosensory mechanism in this species.The main results are as follows:
1.Mating behavior of B. horsfieldi
We observed the mating behavior of B. horsfieldi in the laboratory and confirmed that copulation of this longhorned beetle consisted of three phases:encountering and pair-bonding; mating attempt and ejaculation; post-copulatory guarding. Male showed a dash behavior (rapid approach toward mates) to the female without antennal contact on most occasions (48 of 60 pairs), and then the male mounted the female.Then, he attempted to mate, including licking the middle of the elytra of the female with his maxillary palpi repeatedly, and bending abdomen to contact the tip of the female genitalia. The average mating attempt time was 32.7 min. Only when the female was willing to copulate, ejaculation would be proceed and lasting 61.2 s average time. After ejaculation, 53 of 60 males (88.3%) remained astride the females showing behavior consistent with mate guarding (average time 25.5min).
Five pairs of male and female has been observed for the mating behavior and oviposition behavior during their whole Adult stage. The results showed that the average mating behaviors are 26.2 times.The first peak appeared at 7d-8d after eclosion. The general trend of mating was decreased, but some secondary peaks appeared after then. The peak of mating occurred at 16:00-18:00 daily variation. The average oviposition period of female was about 30 min. During its whole life, the average oviposition rate of a female is 39.8.The average pre-oviposition period of female was 12d, the peak of oviposition occurred in 36d after eclosion.
2. Location and recognition for mate in B. horsfieldi
In olfactometer bioassays, males were significantly attracted to odors from females' volatiles compared with the control air, but were not attracted to volatiles from females with Rosa multiflora Thunb.It is indicating that there are volatiles of females (non-contact sex pheromone) for males to locate but the function of location is weaker than the feeding-host. All of the beetles tested showed no significant different responding in other groups,suggesting that the female was not attracted by the male. Wind tunnel tests showed that sex pheromone was of great importance for orientation of female by male, but the vision effect was limited. Male showed few attractions for female.
The males showed behaviors of mating attempt (licking and abdominal bending) with living females and frozen females.Frozen females, with cuticular hydrocarbons stripped by hexane extraction showed no attraction for tested males, suggesting that males did not recognize females.Reapplying the solvent extract of frozen females to both washed dead males and females caused mating attempts by the tested males, confirming that cuticular hydrocarbons (contact pheromones) played an important role in recognition of females by male B. horsfieldi. The male presented with a glass tube coated with female extract showed the same bending behavior, providing further evidence that contact pheromones play an essential role in mate recognition, and female hexane extract containing female contact pheromones.
Gas chromatography-mass spectrometry analyses of whole-body hexane extract of male and female revealed substantial differences in hydrocarbon profiles, with eight of the compounds in the extracts from females (total 28 compounds) being absent from those of males (total 24 compounds).For limitation of beetle resource and time, we need to do further researches on these compounds to obtain the active components.
3.Cloning of a chemosensory protein (CSP) gene of B. horsfieldi
It was essential of male licking behavior with his maxillary palpi in mate recognition. A full-length BhorCSP2 gene was cloned from maxillary palpi of B. horsfieldi. The open reading-fragment of BhorCSP2 was 384bp, encoding 127 amino acid residues.This deduced amino acid sequence shared some common structural features with chemosensory proteins in insects with four conserved cysteines and a signal peptide with 19 aa in the N-terminal sequence. The amino acids sequence analysis of BhorCSP2 indicated that it was the highest identitiy with that of Tribolium castaneum CSP11 among 30 insect CSP tested.
1.云斑天牛的交配行为
在室内对60对云斑天牛交配行为的观察发现,雄虫有80%是在没有触角接触的情况下与雌虫完成相遇进而抱对的,即雄虫主要不通过触角接触这种方式寻找雌虫；云斑天牛一次完整的交配行为分为三个阶段：相遇抱对、尝试交配和输精、配后保护。从抱对到雄虫输精之前的阶段,统称为尝试交配阶段,该阶段平均需时32.7min,在这期间雄虫有不断舔触雌虫鞘翅的行为；从雄虫开始输精到输精完成平均时间61.2s；88.3%的云斑天牛输精完成后仍然保持着抱对状态即为配后保护阶段,平均用时25.5min。
在室内对云斑天牛整个成虫期交配和产卵行为进行了观察。结果表明平均每对云斑天牛交配26.2次；交配次数第一次高峰出现在羽化出孔后的7d-8d,总的趋势是随着日龄增加交配次数减少；在一天中,交配主要发生在12：00-22：00,特别是16：00-18：00。雌虫从羽化出孔12日后产卵,产卵过程大致持续30min,平均每雌产卵39.8粒,产卵数随着日龄增加而增加,在36日龄产卵量达到峰值,之后减少进入生殖末期。
2.云斑天牛配偶定位和识别
在室内利用“Y”型嗅觉仪测试了云斑天牛雌、雄成虫之间的嗅觉反应。结果表明,以空气为对照时,雌虫对雄虫有明显的吸引作用(P<0.05),而雄虫对雌虫则无明显的吸引作用(P>0.05)；当以补充营养植物野蔷薇为对照时,雄虫对雌虫+野蔷薇、雌虫对雄虫+野蔷薇均无明显的行为趋向(P>0.05)。说明雌虫可能存在非接触性信息素,而雄虫则可能不存在非接触性信息素。
冻死雌虫与活雌虫同样能够引起测试雄虫曲腹的交配反应,但经正己烷浸泡后的雌虫则不会引起雄虫曲腹行为；而将雌虫正己烷浸泡液回滴到正己烷浸泡后的雌虫或雄虫、以及玻璃管上时,均能够引起雄虫的曲腹行为。说明云斑天牛雌虫存在接触性信息素,雄虫通过接触性信息素对雌虫进行识别。
利用气相色谱—质谱联用技术(GC-MS)对云斑天牛雌、雄成虫正己烷提取物进行了分离鉴定。在本试验条件下,云斑天牛雌、雄成虫正己烷提取物共分离鉴定出32种化合物,雌虫28种、雄虫24种；其中有8种化合物是雌虫所特有的,而对于哪种或哪些化合物对雄虫具有行为活性还需要进一步研究。
3.云斑天牛化学感受蛋白基因的克隆和序列分析
云斑天牛雄虫下颚须的舔翅行为在配偶识别方面有重要作用。为了明确其化学感受机制,利用RT-PCR技术克隆获得了云斑天牛下颚须中的化学感受蛋白基因(BhorCSP2),其核苷酸序列可读框全长384bp,编码127个氨基酸残基；推导的氨基酸序列分析表明其有保守的4个半胱氨酸位点和19个氨基酸的信号肽序列,具有昆虫化学感受蛋白的典型特征；通过对BhorCSP2编码的氨基酸和其它昆虫的30种CSP编码的氨基酸进行进化分析,发现该基因与赤拟谷盗TcasCSP11的同源性最高。
Batocera horsfieldi (Hope) (Coleoptera:Cerambycidae) is as a stem borer and a major destructive pest damaging various trees, which causing serious economic and ecology loss in China. It is difficult to control wood-boring pests because of the larvae cryptic feeding habit. With the development of chemecology, control strategies that are based on controlling or killing longhorned beetle adults which are exposed to wild for feeding and copulating. Although some species of cerambycid beetles have been studied, the mechanisms that are involved in behavior and chemical ecology of B. horsfieldi (which belongs to the lamiine) remain unknow. Research on the mating behavior and recognition mechanisms in this species will help in developing integrated pest management tools as well as increasing knowledge of this beetle. Moreover, a chemosensory protein (CSP) in maxillary palpi has also been researched to realize more about the chemosensory mechanism in this species.The main results are as follows:
1.Mating behavior of B. horsfieldi
We observed the mating behavior of B. horsfieldi in the laboratory and confirmed that copulation of this longhorned beetle consisted of three phases:encountering and pair-bonding; mating attempt and ejaculation; post-copulatory guarding. Male showed a dash behavior (rapid approach toward mates) to the female without antennal contact on most occasions (48 of 60 pairs), and then the male mounted the female.Then, he attempted to mate, including licking the middle of the elytra of the female with his maxillary palpi repeatedly, and bending abdomen to contact the tip of the female genitalia. The average mating attempt time was 32.7 min. Only when the female was willing to copulate, ejaculation would be proceed and lasting 61.2 s average time. After ejaculation, 53 of 60 males (88.3%) remained astride the females showing behavior consistent with mate guarding (average time 25.5min).
Five pairs of male and female has been observed for the mating behavior and oviposition behavior during their whole Adult stage. The results showed that the average mating behaviors are 26.2 times.The first peak appeared at 7d-8d after eclosion. The general trend of mating was decreased, but some secondary peaks appeared after then. The peak of mating occurred at 16:00-18:00 daily variation. The average oviposition period of female was about 30 min. During its whole life, the average oviposition rate of a female is 39.8.The average pre-oviposition period of female was 12d, the peak of oviposition occurred in 36d after eclosion.
2. Location and recognition for mate in B. horsfieldi
In olfactometer bioassays, males were significantly attracted to odors from females' volatiles compared with the control air, but were not attracted to volatiles from females with Rosa multiflora Thunb.It is indicating that there are volatiles of females (non-contact sex pheromone) for males to locate but the function of location is weaker than the feeding-host. All of the beetles tested showed no significant different responding in other groups,suggesting that the female was not attracted by the male. Wind tunnel tests showed that sex pheromone was of great importance for orientation of female by male, but the vision effect was limited. Male showed few attractions for female.
The males showed behaviors of mating attempt (licking and abdominal bending) with living females and frozen females.Frozen females, with cuticular hydrocarbons stripped by hexane extraction showed no attraction for tested males, suggesting that males did not recognize females.Reapplying the solvent extract of frozen females to both washed dead males and females caused mating attempts by the tested males, confirming that cuticular hydrocarbons (contact pheromones) played an important role in recognition of females by male B. horsfieldi. The male presented with a glass tube coated with female extract showed the same bending behavior, providing further evidence that contact pheromones play an essential role in mate recognition, and female hexane extract containing female contact pheromones.
Gas chromatography-mass spectrometry analyses of whole-body hexane extract of male and female revealed substantial differences in hydrocarbon profiles, with eight of the compounds in the extracts from females (total 28 compounds) being absent from those of males (total 24 compounds).For limitation of beetle resource and time, we need to do further researches on these compounds to obtain the active components.
3.Cloning of a chemosensory protein (CSP) gene of B. horsfieldi
It was essential of male licking behavior with his maxillary palpi in mate recognition. A full-length BhorCSP2 gene was cloned from maxillary palpi of B. horsfieldi. The open reading-fragment of BhorCSP2 was 384bp, encoding 127 amino acid residues.This deduced amino acid sequence shared some common structural features with chemosensory proteins in insects with four conserved cysteines and a signal peptide with 19 aa in the N-terminal sequence. The amino acids sequence analysis of BhorCSP2 indicated that it was the highest identitiy with that of Tribolium castaneum CSP11 among 30 insect CSP tested.
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