摘要
蜱是一类以吸食哺乳类、鸟类和爬行类血液为生的体外寄生虫，传播的病原体比其它任何节肢类动物都要多，是一种危害严重的疾病媒介。全球每年由蜱类叮咬、传播疾病等引起的损失进二百亿美元，并且过去的二十年中蜱媒疾病的传播范围在逐渐扩大也越来越成为公众健康问题关注的焦点。人类在抵御蜱类寄生方面一直进行着不懈努力。本论文以美洲花蜱Amblyomma americanum和长角血蜱Haemaphysalis longicornis为研究对象，分别从毒理学、生理学、神经生理学和分子生物学方向对植物精油的杀虫活性、蜱中毒后的生理学反应和神经系统电生理活动的变化以及蜱类先天免疫机制进行了研究。论文成果对于蜱类防治药物的研究、更深入的了解杀虫剂导致害虫死亡的最终原因、探索蜱类与病原体相互作用关系和蜱类的适应机制具有重要意义。本论文主要的研究结果如下：
1、以饥饿阶段美洲花蜱若蜱为对象，利用毒理学方法检测了墨西哥牛至Lippia graveolens、多香果Pimenta dioica和荆芥Nepeta cataria L.三种植物粗提精油的杀虫活性。结果显示，三种植物精油对美洲花蜱都具有致死毒性，尤其在处理后24小时死亡率达到80%以上。半数致死剂量（LC50）以及处理后3小时和6小时死亡率和剂量之间的线性关系显示，同时间点墨西哥牛至的毒性显著高于多香果和荆芥，而后两者未表现出显著差异。
为了为后面工作提供依据，对两种化学性杀虫剂—氯菊酯permethrin和阿米曲拉amitraz作用于美洲花蜱雌蜱的致死和半致死剂量也进行了测试，两者的致死和半致死剂量分别为0.010%、0.005%和16μg μl~(-1)、8μg μl~(-1)。
2、利用直流呼吸运动计量法对氯菊酯和阿米曲拉以及两者混合剂作用后美洲花蜱雌蜱的气体交换和水分平衡进行了检测。结果显示，所有对照组个体的呼吸都为不连续气体交换模式。氯菊酯处理的个体最初呼吸强烈之后转变为高频率的不连续模式，而阿米曲拉和混合剂处理后的呼吸呈明显的混乱状态。正常组和丙酮处理组的标准代谢率分别为0.461and0.452μl h~(-1)，且两组间无显著差异。三个杀虫剂处理组的标准代谢率显著高于对照组，氯菊酯处理组为1.054μl h~(-1)、阿米曲拉处理组为1.392μl h~(-1)、混合剂处理组为1.520μl h~(-1)。氯菊酯处理后1-2小时内虫体大量失水，且在不连续呼吸模式出现后每次呼吸都伴随水分丢失。与氯菊酯处理组相比，混合剂处理后个体同样大量失水，但这种失水现象不局限于处理后的1-2小时而是整个记录过程，然而未发现与呼吸同步的失水。对照组和阿米曲拉处理组未表现大量失水的现象。这些结果表明氯菊酯破坏了蜱体内水分保持机制，阿米曲拉使气门控制系统过度兴奋，而混合剂则引起两种药物的双重反应。
3、采用电生理学手段研究了氯菊酯、阿米曲拉及其混合剂处理后美洲花蜱附肢运动肌电信号活动的变化。实验中共记录到两种信号，分别命名为S1和S2。正常情况下蜱附肢中的神经信号具有较明显的模式，具有清晰的S1放电簇且记录到的两种信号差异明显。氯菊酯、阿米曲拉以及两者的混合剂处理后运动肌电信号出现了不同的变化。氯菊酯处理后S1放电簇频率降低但时长延长，阿米曲拉处理后S1放电簇逐渐紊乱，最终无明显规律，混合剂处理后出现了叠加性的反应，而处理1小时之后的反应也显示双重机制作用下虫体神经系统功能瘫痪。
4、从长角血蜱雄蜱附腺cDNA文库中分离得到编码抗菌肽的基因。基因全长349bp，编码79个氨基酸的防御素蛋白，基因被命名为HlMS-defensin。反转录PCR显示HlMS-defensin只在雄蜱中表达。采用real-time PCR技术分析了雄蜱不同组织和各吸血阶段HlMS-defensin的表达情况。结果表明HlMS-defensin主要在雄蜱附腺中表达并且随着吸血时间的延长表达量也不断升高。对成熟肽合成蛋白的抗菌活性分析显示HlMS-defensin对革兰氏阳性菌、革兰氏阴性菌和真菌都具有活性。上述结果表明蜱类很可能通过抗菌肽来保护生殖系统不被病原微生物感染。
Ticks are obligate ectoparasite living by blood of mammals, birds and reptilia,transmit more pathogenic microorganisms than any other arthropod vectors, and areamong the most important vectors of pathogens. On global basis, economic lostcaused by tick and tick-borne diseases is almost billions of dollars. In the post twodecades, tick-borne diseases had increased incidence of and are becoming one of thefocus of public health problem. Unremitting work is making against ticks. In thecurrent study, toxicological, physiological, neurophysiological and molecularbiological research, using Amblyomma americanum and Haemaphysalis longicornis,was made to (1) test the anti-tick activities of three essential oils,(2) investigatephysiological response of the poisoned ticks and the effects of chemical pesticideson neurophysiology and (3) explore the innate immunity of ticks. The results will besignificant for acaricidal research, further understand the ultimate causes of pesticideintoxication and explore the interaction of tick and pathogens and the adaptationmechanism of ticks. Major results of the current study were as followed:
1. Toxicological efficacy of essential oils from Lippia graveolens, Pimentadioica, Nepeta cataria L. was tested against A. americanum nymphs. All the threeessential oils showed lethal toxicity. Especially after24h of treatment, highmortalities were observed. LC50and the relationship between dose and mortality of3and6h post-treatment revealed that toxicity of Lippia graveolens was significantlyhigh than Pimenta dioica and Nepeta cataria L. on same time point, toxicity of thelatter two didn't differ significant.
In order to provide basis for subsequent work, the lethal and sublethal dose oftwo chemical pesticides, permethrin and amitraz, against A. americanum females,were also toxicologically tested. The lethal and sublethal doses were0.010%、0.005%and16μg μl~(-1)、8μg μl~(-1)for permethrin and amitraz, respectively.
2. Changes in the gas exchange and water balance of lone star tick, A.americanum poisoned by permethrin and amitraz, were examined using flow-through respirometry. All control ticks treated by nothing or acetone exhibiteddistinct discontinuous gas exchange cycle (DGC). The pattern of permethrin-treatedticks was characterized by impetuous ventilation followed by highly frequent DGC;while the ventilation of amitraz and permethrin+amitraz (mixture)-treated ticksremained apparently chaotic. Standard metabolic rates of untreated andacetone-treated ticks were0.461and0.452μl h~(-1), respectively, and didn't differsignificantly from each other (p>0.05). Acaricides-treated ticks exhibitedsignificant increasing in SMR (p <0.05;1.054,1.392and1.520μl h~(-1)for permethrin,amitraz and mixture-treated ticks, respectively). Permethrin also induced obviouswater loss in ticks. A mass of water lost in the first1-2h post-treatment wasobserved and coincidence between water loss and CO2emission was foundespecially during DGC periods. In contrast to permethrin-treated ticks, no matchbetween water loss and gas exchange was found in mixture-treated ticks, but greatwater loss was demonstrated throughout the recordings. No obvious water loss wasobserved in control and amitraz-treated ticks. The results suggested that permethrinupset the water maintenance mechanisms of ticks; amitraz overexcited the spiracularcontrolling mechanisms; and thereby, the mixture of permethrin and amitraz resultedin chaotic ventilation and great water loss.
3. The activity of leg motor of A. americanum, before and after treated bypermethrin, amitraz and their mixture, was recorded via electrophysiologicaltechnique. Two kinds of spike were detected and were termed as S1and S2. Normalticks show distinct pattern, S1burst was clear and the two kinds of spike differedobviously. Various signals were detected in permethrin, amitraz and mixture treatedticks. Permethrin-treated ticks exhibited frequent and prolonged burst, the burst ofamitraz-treated ticks became disordered generally after treatment and lost the patternfinally. Mixture-treated ticks showed both response of permethrin and amitraz singletreated ticks and the response of1h post-treatment suggested that the nervous systemwas paralyzed.
4. A novel gene was identified from a cDNA library of the male accessoryglands of Haemaphysalis longicornis. The full-length cDNA of the gene was349bp, encoding a79-amino acid defensin-like protein and therefore the protein was namedas HlMS-defensin. Reverse transcriptase-PCR results suggested that this gene wasexpressed exclusively in male ticks. The tissue expression pattern and the mRNAlevels of HlMS-defensin during blood feeding were determined using real-time PCR.HlMS-defensin was expressed predominately in the male accessory gland and wasup-regulated during blood feeding. The antimicrobial activity of a synthetic peptidebased on the predicted mature portion of HlMS-defensin was examined against avariety of Gram-positive and Gram-negative bacteria and fungi. It appears that ticksuse the antimicrobial peptide to protect their reproductive tracts from microbialinfections.
引文
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