南亚果实蝇种群特征及其对食料和热胁迫的生理调控机制
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摘要
南亚果实蝇Bactrocera tau Walker属双翅目Diptera，实蝇科Tephritidae，果实蝇属Bactrocera Macquart，是危害黄瓜Cucumis Sativus L．、南瓜Cucurbita Moschala L．、丝瓜Luffa Cylindrical L．、冬瓜Benincasa hispida(Thunb)Coqn和苦瓜Momordica Charantia L．等果蔬作物的一种重要害虫。近年来，在我国广东、福建以及云南、四川和重庆等省市发生，危害程度呈加重的趋势，严重影响果蔬作物的生产。尽管南亚果实蝇在果蔬作物生产过程中具有重要经济意义，然而其成灾及适应环境的机理尚不明确。据此，本论文立足于国内外的研究现状，借助昆虫生态、昆虫生理、生物化学以及生物统计等学科知识，在研究南亚果实蝇种群特征的基础上，探讨种群适应食料和温度的生化机理。本研究得到教育部博士点基金项目(20050625005)以及重庆市科委攻关项目(7468)的资助，经过近3年的努力，取得以下研究结果。
1重庆实蝇类害虫的种类及地理分布
2005-2006年，采用诱蝇醚、诱蝇酮和水解蛋白等5种实蝇诱集剂，对重庆市30多个区县的实蝇类害虫进行全面诱集，共收集、整理和鉴定实蝇标本174879头。分属以下8个种内，即南亚果实蝇Bactrocera tau Walker、黑颜果实蝇B．diaphora Hendel、宽带果实蝇B．scutellata Hendel、桔大实蝇B．minax Enderlein、瓜实蝇B．cucurbitae Coquillett、桔小实蝇B．dorsalis Hendel、普通果实蝇B．caudate Fabricius和三点棍腹实蝇Dacus trimacula。其中，桔小实蝇、瓜实蝇和三点棍腹实蝇在重庆地区属首次发现。
采用优势度指数分析了重庆地区各类实蝇害虫的优势度。2005年，南亚果实蝇、黑颜果实蝇、宽带果实蝇、桔大实蝇、瓜实蝇、桔小实蝇和普通果实蝇的优势度指数分别为0.8225，0.1333，0.0376，0.0055，0.0009，0.0001和0.0001；2006年，以上7种实蝇和三点棍腹实蝇的优势度指数则分别为0.7651，0.0889，0.1304，0.0003，0.0028，0.0104，0.0010和0.0011。表明南亚果实蝇是重庆地区的优势实蝇种群；黑颜果实蝇和宽带果实蝇次之；其余实蝇种群数量较低。
南亚果实蝇、黑颜果实蝇和宽带果实蝇分布于重庆设置诱集点的各个区县。桔大实蝇、普通果实蝇和三点棍腹实蝇分布于重庆局部区域。2005年诱集到桔小实蝇9头，分布于江北、永川、垫江和涪陵4个区县；2006年诱集到95头，分布于江北、九龙坡等16个区县，分布范围扩大。2005年诱集到瓜实蝇10头，2006年诱集到956头，分布于巴南、九龙坡、秀山、合川和长寿5个区县，诱集数量增加，分布区域不变。
2南亚果实蝇的为害特性及风险分析
对南亚果实蝇种群的危害特性及风险性进行了系统评价。在重庆地区，南亚果实蝇幼虫主要危害黄瓜、南瓜、丝瓜、冬瓜和苦瓜等多种果蔬作物，导致果实腐烂，提前脱落，在受害区域常造成严重的经济损失。以上5种作物中，该虫对南瓜和丝瓜的危害重，黄瓜次之，苦瓜轻。根据有害生物危险性评价指标体系和有害生物的危险性综合评价标准，对该虫的风险性进行了综合评价，发现南亚果实蝇的危险性R值为1.96，属于中度危险的有害生物。
3南亚果实蝇种群特征及其与环境湿度的关系
通过室内外实验，对南亚果实蝇的行为特性、结构特征以及环境湿度对该种群的影响等方面进行了研究。在重庆地区，南亚果实蝇1年发生3-5代，世代重叠明显。成虫集中在近地面范围内活动，具有多次交尾习性。幼虫取食果肉，破坏果实组织，老熟后钻出果实，入土化蛹。70％以上的老熟幼虫在0-5cm厚的土层中化蛹，存活率达88％以上；3.74％老熟幼虫的化蛹深度超过10cm，存活率低于65％；当土壤深度超过25cm时，仅3.33％的个体能够存活。蛹在一天内各个时段均可羽化，但以每天上午6：00-10：00羽化的数量较多，占全天羽化量的85.80％。
南亚果实蝇对黄瓜、南瓜、冬瓜、丝瓜、苦瓜和柑桔幼果具有不同的取食和产卵选择性。在黄瓜、南瓜和丝瓜上，成虫的落虫数量分别为25.33，21.33和22.67头，产卵量分别为98.00，86.00和103.33粒，均显著高于其余3种果实上的平均落虫量和产卵量。幼虫在黄瓜、南瓜和丝瓜上取食的数量亦较高，分别占幼虫总量的23.33％，31.44％和21.67％；而在柑桔上仅占幼虫总量的2.22％。成虫和幼虫均对受害与未受害果实的选择特性表现出一定的差异，在受害果实上的虫口数量较多，未受害果实上的虫口数量较少。因此，在这6种寄主植物中，南亚果实蝇喜好选择在黄瓜、南瓜和丝瓜以及受害的果实上取食和产卵。
土壤相对水分含量(SRWC)以及空气相对湿度(RH)对南亚果实蝇蛹的生长发育和失水动态均有影响。占78.83％的老熟幼虫集中在SRWC 20-60％的土壤中化蛹。当SRWC为100％时，蛹不能羽化；当SRWC为40％和60％时，蛹具有较短的发育历期(7.54d和7.65d)和较高的羽化率(83.58％和93.3％)。在0％，52％，76％和100％这4个RHs下，蛹的水分损失率随着处理时间的延长而增加，7日蛹水分损失率分别达到63.521％(RH0％)，45.42％(RH52％)，34.23％(RH76％)和10.99％(RH100％)。在这4个RHs下，蛹的发育历期和羽化率亦存在显著差异，在RH100％时，具有较短的发育历期(8.07d)和较高的存活率(90.55％)；而RH0％时，仅3.03％的蛹能够羽化。因此，SRWC 100％和RH0％均不利于南亚果实蝇蛹的发育和存活。
4不同食料对南亚果实蝇生长发育及酶学特性的影响
在室内温度28±0.5℃、相对湿度RH 70±5％、光周期14 L：10 D条件下，研究了南亚果实蝇分别取食黄瓜、南瓜、丝瓜、苦瓜、柑桔和人工饲料时的生长发育特性和繁殖能力。结果表明，南亚果实蝇取食以上各种食料后，幼虫的发育历期和存活率均存在显著差异，分别为4.58d，5.08d，4.50d，6.15d，5.38d，5.72d和86.83％，83.39％，81.19％，51.17％，45.67％，30.33％；成虫的产卵量分别为99.56，84.67，85.91，33.97，8.24和3.87粒，表明取食黄瓜、南瓜和丝瓜后成虫的产卵量较高，取食人工饲料的较低。南亚果实蝇分别取食以上6种食料后，内禀增长率分别为0.0844，0.0572，0.0601，0.0604，-0.0014和-0.0159；种群的趋势指数分别为31.69，20.78，22.62，3.52，0.54和0.11。以上结果均表明，黄瓜、南瓜和丝瓜是该种群生长发育的适宜食料。
超氧化物歧化酶(SOD)、过氧化物酶(POD)和过氧化氢酶(CAT)是生物体内清除活性氧自由基的多酶复合体系，测定了南亚果实蝇分别取食黄瓜、南瓜、冬瓜、丝瓜和苦瓜后，幼虫、蛹和成虫体内SOD、POD和CAT的活性。结果表明，南亚果实蝇幼虫、蛹和成虫体内SOD、POD和CAT的活性均以取食苦瓜时较高，取食冬瓜时次之，取食黄瓜和南瓜时则较低。SOD的活性以幼虫阶段较高，成虫阶段较低；POD和CAT的活性均以成虫阶段较高，蛹阶段较低。表明南亚果实蝇幼虫取食适宜的食料，虫体抗氧化酶的活性较低；而取食相对不适合的食料时，抗氧化酶的活性提高。
羧酸酯酶(CarE)、酸性磷酸酯酶(ACP)、碱性磷酸酯酶(ALP)、细胞色素氧化酶(P450s)和谷胱甘肽S-转移酶(GST)是生物体内降解内源和外源性有毒化合物的重要解毒酶，研究了5种食料对南亚果实蝇这些解毒酶活性的影响。双因子方差分析表明，南亚果实蝇取食黄瓜、南瓜、丝瓜、冬瓜和苦瓜后，解毒酶活性存在显著差异。其中，羧酸酯酶(CarE)活性在黄瓜和南瓜上较高，细胞色素氧化酶P450 O-脱甲基和谷胱甘肽S-转移酶(GST)活性在苦瓜上较高，酸性磷酸酯酶(ACP)和碱性磷酸酯酶(ALP)活性却分别在黄瓜和南瓜上较低。幼虫、蛹和成虫解毒酶的活性亦存在显著差异，成虫具有较高的CarE活性；幼虫具有较高的细胞色素P450 O-脱甲基、GST和ALP活性，但具有较低的ACP活性：除ACP外，蛹的解毒酶活性均较低。因此，南亚果实蝇解毒酶活性受寄主果实种类以及该种群本身发育阶段的影响。
多酚氧化酶是昆虫生命活动过程中的重要调节酶，研究了南亚果实蝇以南瓜为食料时，不同虫态多酚氧化酶的活性和性质。以邻苯二酚为底物，南亚果实蝇不同虫态多酚氧化酶活力大小依次为：3龄幼虫＞成虫＞2龄幼虫＞1龄幼虫＞蛹。在pH6.5，温度34-37℃时的活性较高；在40℃以上时，随着温度的升高酶活性下降。2龄幼虫的K_m和V_(max)较大，表明多酚氧化酶对底物的亲和力较弱，催化能力较强。而以L-多巴为底物时，成虫K_m和3龄幼虫V_(max)较大，表明成虫多酚氧化酶对底物的亲和力较弱，3龄幼虫多酚氧化酶对底物的催化能力较强。因此，南亚果实蝇体内多酚氧化酶的活性和性质与个体的发育阶段密切相关。
5温度对南亚果实蝇生长发育及能源物质含量的影响
以南瓜为食料，研究了6个温度梯度对南亚果实蝇种群生长发育和繁殖的影响。从19℃至28℃，南亚果实蝇幼虫的发育历期随温度的升高而缩短，存活率则随温度的升高而增加；当温度高于28℃时，发育历期随温度的升高而延长，存活率则降低。从19℃至32℃，成虫寿命均随温度的升高而缩短，产卵量在28℃时较高。生殖力生命表参数和种群趋势指数均表明28℃时是南亚果实蝇生长发育和繁殖的适宜温度。
探讨了42℃高温胁迫不同时间对南亚果实蝇成虫寿命以及虫体能源物质即多糖、甘油三酯和可溶性蛋白含量的影响。结果表明，连续处理10d，以每天胁迫10min时成虫寿命较长；连续处理15d时，则以每天胁迫5min时成虫寿命较长。高温胁迫导致虫体多糖贮存量下降，连续处理15d，每天分别胁迫为5，15，30和60min后，多糖含量的损失百分率分别达到为2.04％，2.08％，5.04％和5.93％。对于甘油三酯，当胁迫时间为5min时，虫体甘油三酯的含量较低，此后随着胁迫时间的延长，虫体甘油三酯含量的累积却增加。高温胁迫也会促进南亚果实蝇对蛋白质的利用，随着胁迫时间的延长，虫体蛋白质的贮存量下降。以上结果可以推出，高温胁迫将提高南亚果实蝇虫体对多糖和蛋白质的消耗，而增加甘油三酯的累积。
本文研究了重庆地区南亚果实蝇优势种群的生物生态学特性以及环境因子胁迫后体内多种化学物质含量和活性的变化规律，揭示了南亚果实蝇在重庆地区的发生规律、成灾机理以及适应环境食料和温度的生化机理。研究结果不仅为南亚果实蝇的发生预测和综合治理提供必要的科学依据，亦为南亚果实蝇的分子生物学研究奠定重要的理论基础。
The fruit fly Bactrocera tau Walker, belonging to the order Diptera and the family Tephritidae, mainly infests Cucumis L., Cucurbita L., Luffa L., Benincasa Savi, Momordica L. and other fruits and vegetables. It has taken place in tropic and subtropic regions such as China, Japan, Thailand, Malaysia, Laos, Philippines, Cambodia, India, Bhutan, Indonesia, Sri Lanka and has caused enormous economic losses to fruit and vegetable growers. In China, the fruit fly caused Siraitia grosvenorii Swingle damage up to 21-34% in Guangxi province in 1990 and was injurious to various types of vegetables in Chongqing district in 2004. Due to its economic importance, the study on biological and biochemical characters of this fruit fly was very necessary in the course of the production of fruit and vegetable crops. Based on the research status, the study on the population biological and biochemical characters, especially the physiological mechanisms of the fruit fly adaptable to diets and environmental temperature was carried out systematically. This study was supported by the Specialized Research Fund for the Doctoral Program of Higher Education (20050625005) and the Basic Research Foundation of Chongqing Science Committee (7468), and lasted almost 3 years. The main results are summarized as follows.
1 The species richness and geographical distribution of fruit flies in Chongqing
By the Regional Fruit Fly Project in 2005-2006, systematic trapping using methyl eugenol, cue-lure and hydrolytic protein had been conducted in almost 30 regions of Chongqing. The total 174879 fruit fly samples trapped had confirmed the presence of Bactrocera tau Walker, B. diaphora Hendel, B. scutellata Hendel, B. caudate Fabricius, B. minax Enderlein, B. cucurbitae Coquillett, B. dorsalis Hendel and Dacus trimacula. Among them, B. cucurbitae, B. dorsalis and Dacus trimacula was firstly found in Chongqing district.
Species relative abundance of fruit flies in Chongqing region was analyzed by dominance indexes. The dominance indexes of these fruit flies varied greatly among the 8 kinds of species. In 2005, the dominance indexes of B. tau, B. diaphora, B. scutellata, B. caudate, B. minax, B. cucurbitae and B. dorsalis were 0.8225, 0.1333, 0.0376, 0.0055, 0.0009, 0.0001 and 0.0001, respectively. In 2006, they were 0.7651, 0.0889, 0.1304, 0.0003, 0.0028, 0.0104, 0.0010 and 0.0011 (Dacus trimacula), respectively. The result indicated that B. tau was the dominant population in Chongqing region, B. diaphora and B. scutellata were the second great population, and the others were relatively low in quantities.
According to the identification result of samples collected from different regions of Chongqing, they showed that B. tau, B. diaphora and B. scutellata widely distributed in Chongqing district, but B. minax, B. caudate and D. trimacula were found just in several locations. For B. dorsalis, 9 individuals were captured and distributed in Jiangbei, Yongchuan, Xiushan, Dianjiang and Fuling in 2005, and 95 individuals were captured and distributed in 16 regions including Jiangbei, Jiulongpo, Beibei, and so on in 2006. It showed that the distribution range obviously expanded in 2006. For B. cucurbitae, 10 individuals were trapped in 2005 and 956 individuals were trapped in 2006, and it indicated that the quantity captured in 2006 was significantly higher than that in 2005.
2 The damage character and risk analysis of B. tau population
By the field investigation and the previous study, we comprehensively evaluated the damage character and risk of B. tau. The result showed that B. tau mainly infected cucumber, pumpkin, towel gourd, white gourd, balsam pear and other fruits and vegetables in Chongqing district. Host surveys for this fruit fly in field had confirmed that the damage rate was highest on pumpkin and towel gourd, then on cucumber, and lowest on balsam pear among the five kinds of host fruits above-mentioned. Based on the system and indices of pest risk analysis, we estimated the risk of this fruit fly, and found that the index value of pest risk analysis (R) attained to 1.96. From the result, it could be inferred that this fruit fly was dangerous to some fruits and vegetables, and had posed a threat to other regions in China.
3 The population character of B. tau and its relation with environmental humidity
B. tau had 3-5 generations a year and the generations overlapped in Chongqing district. It was divided into 3 instars in larval stages, and overwintered as pupae in loose soils. Generally, adults mated for several times in the whole life, and the female laid its eggs in clutches under the skin of fruits. The larva or maggot fed on the flesh of fruits. In the process, bacteria were introduced into the fruit causing the fruit to break down or rot. More than 70% mature larvae pupated in the soil with 0-5cm depth, and the survival rate of pupae was higher than 88%. There were 3.74% individuals digging into the soil with 10-25cm depth, and the survival rate was lower than 65%. Only 3.33% individuals could survive in the soil depth over 25cm. The adult could emerge at every time of a day, but mostly aggregated on the time 6: 00-10: 00.
The taxis response and selective propensity of B. tau to six types of fruits cucumber, pumpkin, towel gourd, white gourd, balsam pear and orange were evaluated under indoor experimental conditions. The result demonstrated that most of the adults would obviously prefer to aggregate on cucumber, pumpkin and towel gourd to gain nutrition or lay eggs. Among them, 23.68%, 19.94% and 21.18% of individuals gathered to cucumber, pumpkin and towel gourd, and 21.57%, 18.93% and 22.74% of eggs were lay on the same three host fruits, respectively. It was suitable, adopting linear regress equation, to describe the relationship between the mean quantities and the mean oviposition quantities. As adults, most larvae also aggregated on cucumber (23.33%), pumpkin (31.44%) and towel gourd (21.67%), but little individuals aggregated on orange (2.22%). In addition, the taxis response of the fruit fly to damaged and undamaged fruits was also different in the six kinds of host fruits, and＞50% of individuals would gather to the damaged fruits. From these results, it could seen that the fruit fly preferred to aggregate on cucumber, pumpkin, towel gourd and damaged fruits, but little on orange to gain nutrition or lay eggs.
The effect of soil relative water content (SRWC) and air relative humidity (RH) on the pupal development and survival and the water loss dynamics of B. tau were evaluated under indoor experimental conditions. The pupating selectivity of mature larvae, the pupal duration and eclosion rate varied significantly in the soils with SRWC 0%, 20%, 40%, 60%, 80% and 100%. Most of the mature larvae aggregated on the soils with SRWC 20-60% although they could become pupae in the soils with different water content. When SRWC attained to 100%, all of the pupae failed to survive; SRWC 40% and 60% were most suitable for the pupal development, with the shortest pupal duration (7.54d and 7.65 d) and the highest eclosion rate (83.58% and 93.3%). In the 4 air RHs of 0%, 52%, 76% and 100%, the pupal water loss rates increased with the increase of the treatment time, and reached up to 63.521% for RH 0%, 45.42% for RH 52%, 34.23% for RH 76% and 10.99% for RH 100% at the last day before eclosion. It was suitable, adopting the quadratic equations, to describe the relationship between the water loss rate of pupae accumulated by day and pupal age in day. The pupal duration and eclosion rate were also significantly different in the 4 air RHs. The shortest duration (8.07d) and the highest survival rate (90.55%) of pupae were attained in the air RH 100%, and little individuals could survive in the air RH 0%. The results implied that high soil water content and low air humidity were unfit for the development and survival of B. tau pupae.
4 The influence of diets on the growth and enzyme characterization of B. tau
The effect of different diets including cucumber, pumpkin, towel gourd, balsam pear, orange and artificial diet on the growth and fecundity of B. tau were evaluated at 28±0.5℃, RH 70±5% and photoperiod of 14:10 hours (L: D). The result demonstrated that the developmental duration and survival rate of larvae and the reproduction rate of adults differed significantly among the 6 kinds of diets. When the larva were reared on the above diets, the larval durations were 4.58d, 5.08d, 4.50d, 6.15d, 5.38d, 5.72d, the survival rates were 86.83%, 83.39%, 81.19%, 51.17%, 45.67%, 30.33%, and the total reproduction quantities per female were 99.56, 84.67, 85.91, 33.97 and 8.24, respectively. The result suggested that the larval durations were shorter, the survival rates were higher and the reproduction quantities per female were also higher on cucumber, pumpkin and towel gourd. According to the life-table parameters and the population trend indexes, it could be concluded that cucumber, pumpkin and towel gourd were the most suitable diets for the development and fecundity of this fruit fly, but contrary for orange and artificial diet.
Superoxide dismutase (SOD), Peroxidase (POD) and Catalase (CAT) are considered to be an important physiological and biochemical mechanism of organisms against adverse environmental factors. The soluble protein contents and those enzyme activities in B. tau fed on cucumber, pumpkin, towel gourd, white gourd and balsam pear were determined by biochemical methods. The result indicated that the protein contents and those detoxification enzyme activities varied significantly when the fruit fly were reared on different diets—cucumber, pumpkin, towel gourd, white gourd and balsam pear. Among the 5 kinds of diets, the protein contents in this fruit fly were highest on balsam pear while lowest on cucumber and white gourd. The SOD, POD and CAT activities were highest on balsam pear, then white gourd, but lowest on cucumber and pumpkin, and those enzyme activities also varied greatly in different developmental stages of this fruit fly. The SOD activity was highest at the larval stage, and lowest at the adult stage; the POD and CAT activities were highest at the adult stage, but lowest at pupal stage. Based on the above study, it indicated that the antioxidant activity in B. tau was lower when the fruit fly was reared on suitable diets, than that on unsuitable diets.
This study was conducted to determine whether the activity of detoxification enzymes, such as carboxylesterase (CarE), acid phosphatase (ACP), alkaline phosphatase (ALP), cytochrome P450-dependent O-demethylase and glutathione S-transferase (GST) in Bactrocera tau (Walker), respond to different diets and developmental stages of this fruit fly. Two-way ANOVA showed that protein contents and these detoxification enzyme activities varied significantly when the fruit fly were reared on different diets cucumber, pumpkin, towel gourd, white gourd and balsam pear. Among the five kinds of diets, protein contents in the fruit fly were highest on balsam pear while lowest on cucumber and white gourd. The CarE activity was highest on cucumber and pumpkin, cytochrome P450-dependent O-demethylase and GST activities were highest on balsam pear, but ACP and ALP activities were lowest on cucumber and pumpkin, respectively. In addition, these detoxification enzyme activities were also significantly different in the larval, pupal and adult stages of the fruit fly. Adults had the highest CarE activity, and larvae had the highest ALP, cytochrome P450-dependent O-demethylase and GST activities, but the lowest ACP activity. Generally, these detoxification enzyme activities in the pupal stage were lower than those in the larval and adult stages except for ACP. The result indicated that these detoxification enzyme activities of B. tau were closely related to host plant species and developmental stages of this fruit fly.
Polyphenoloxidase (PPO), a group of copper proteins thatare widely distributed from bacteria to mammals, catalyzes the oxidation of hydroxyphenols to their quinone derivatives, which then spontaneously polymerize. The kinetic properties of PPO from B. tau different stages were compared and determined by a series of chemical methods. The result showed that PPO activity differed significantly at different stages in this insect. The PPO activity of the 3rd instar was highest and that of pupae was lowest among the 1st instar, the 2nd instar, the 3rd instar, pupae and adults. The order of the enzyme activity was the 3 instar＞adults＞the 2nd instar＞the 1st instar＞pupae. The optimum pH was 7.0 and the best temperature was 34-37℃for the tested PPO. The kinetic parameter for the oxidation of catechol by PPO from the different stages was determined and compared, and it showed that the affinity of PPO to the substrates catechol was higher, but the catalytic activity was lower for the 2nd instar. At the same time, for the substrates L-DOPA, the K_m for adults and V_(max) for the 3rd instar of PPO were highest, it showed that the affinity of PPO for adults was lower, and the catalytic activity for the 3rd instar was higher than the other stages. From the above conclusion, it could be inferred that the property of PPO activities is closely associated with the developmental stages of B. tau.
5 The influence of temperature on the growth and energy resources of B. tau
The growth and fecundity of B. tau fed on pumpkin were investigated at 6 constant temperatures 19℃, 22℃, 25℃, 28℃, 31℃and 34℃in a series of indoor trials. ANOVA showed that temperature affected significantly the developmental duration and reproduction capacity. The developmental duration of larvae varied from 6.31d at 28℃to 10.09d at 19℃, and the survival rate varied from 27.03% at 34℃to 74.76% at 28℃. The female adult at 19℃and 22℃had significantly greater longevity (96.29d and 94.00d) than those at the other temperature, and had the greatest reproduction capacity (105.91 eggs per female) at 28℃. Accordingly, the intrinsic rate of increase (r_m) and the population trend index were highest (0.0572 and 17.28, respectively) for this fruit fly at 28℃. The results implied that the temperature of 28℃were the most suitable for the population development and fecundity.
The longevity and energy sources including polysaccharides, triacylglycerol and soluble protein of B. tau submitted to heat shocks (42℃) of various durations (5,15, 30 or 60 min daily) for 10d or 15d were determined systematically by ecological and biochemical methods. A slight longevity increase was observed in 15min daily for 10d and in 5min daily for 15d, but longer shocks had negative effects. Fruit flies submitted to the procedure providing a longevity increase did not show a higher polysaccharide and protein contents than those in the other treatment durations. However, the highest polysaccharide and protein contents were observed in the control flies. Except for the control flies, the accumulation of triacylglycerol of this fruit fly increased responding to the treated durations. The above results suggested that under the heat shocks conditions, polysaccharides and soluble protein were the main energy sources, while the utilization of triacylglycerol decreased.
Generally, the study on the bio-ecological characteristics, and the influence of diets and heat shock on the physiological and biochemical properties of B. tau has clearly elucidated the occurrence regulation, formative mechanism and biochemical mechanism of this fruit fly suitable to diets and temperature. The result will provide necessary information for IPM program, and enrich the whole theoretical system of fruit flies.
1重庆实蝇类害虫的种类及地理分布
2005-2006年，采用诱蝇醚、诱蝇酮和水解蛋白等5种实蝇诱集剂，对重庆市30多个区县的实蝇类害虫进行全面诱集，共收集、整理和鉴定实蝇标本174879头。分属以下8个种内，即南亚果实蝇Bactrocera tau Walker、黑颜果实蝇B．diaphora Hendel、宽带果实蝇B．scutellata Hendel、桔大实蝇B．minax Enderlein、瓜实蝇B．cucurbitae Coquillett、桔小实蝇B．dorsalis Hendel、普通果实蝇B．caudate Fabricius和三点棍腹实蝇Dacus trimacula。其中，桔小实蝇、瓜实蝇和三点棍腹实蝇在重庆地区属首次发现。
采用优势度指数分析了重庆地区各类实蝇害虫的优势度。2005年，南亚果实蝇、黑颜果实蝇、宽带果实蝇、桔大实蝇、瓜实蝇、桔小实蝇和普通果实蝇的优势度指数分别为0.8225，0.1333，0.0376，0.0055，0.0009，0.0001和0.0001；2006年，以上7种实蝇和三点棍腹实蝇的优势度指数则分别为0.7651，0.0889，0.1304，0.0003，0.0028，0.0104，0.0010和0.0011。表明南亚果实蝇是重庆地区的优势实蝇种群；黑颜果实蝇和宽带果实蝇次之；其余实蝇种群数量较低。
南亚果实蝇、黑颜果实蝇和宽带果实蝇分布于重庆设置诱集点的各个区县。桔大实蝇、普通果实蝇和三点棍腹实蝇分布于重庆局部区域。2005年诱集到桔小实蝇9头，分布于江北、永川、垫江和涪陵4个区县；2006年诱集到95头，分布于江北、九龙坡等16个区县，分布范围扩大。2005年诱集到瓜实蝇10头，2006年诱集到956头，分布于巴南、九龙坡、秀山、合川和长寿5个区县，诱集数量增加，分布区域不变。
2南亚果实蝇的为害特性及风险分析
对南亚果实蝇种群的危害特性及风险性进行了系统评价。在重庆地区，南亚果实蝇幼虫主要危害黄瓜、南瓜、丝瓜、冬瓜和苦瓜等多种果蔬作物，导致果实腐烂，提前脱落，在受害区域常造成严重的经济损失。以上5种作物中，该虫对南瓜和丝瓜的危害重，黄瓜次之，苦瓜轻。根据有害生物危险性评价指标体系和有害生物的危险性综合评价标准，对该虫的风险性进行了综合评价，发现南亚果实蝇的危险性R值为1.96，属于中度危险的有害生物。
3南亚果实蝇种群特征及其与环境湿度的关系
通过室内外实验，对南亚果实蝇的行为特性、结构特征以及环境湿度对该种群的影响等方面进行了研究。在重庆地区，南亚果实蝇1年发生3-5代，世代重叠明显。成虫集中在近地面范围内活动，具有多次交尾习性。幼虫取食果肉，破坏果实组织，老熟后钻出果实，入土化蛹。70％以上的老熟幼虫在0-5cm厚的土层中化蛹，存活率达88％以上；3.74％老熟幼虫的化蛹深度超过10cm，存活率低于65％；当土壤深度超过25cm时，仅3.33％的个体能够存活。蛹在一天内各个时段均可羽化，但以每天上午6：00-10：00羽化的数量较多，占全天羽化量的85.80％。
南亚果实蝇对黄瓜、南瓜、冬瓜、丝瓜、苦瓜和柑桔幼果具有不同的取食和产卵选择性。在黄瓜、南瓜和丝瓜上，成虫的落虫数量分别为25.33，21.33和22.67头，产卵量分别为98.00，86.00和103.33粒，均显著高于其余3种果实上的平均落虫量和产卵量。幼虫在黄瓜、南瓜和丝瓜上取食的数量亦较高，分别占幼虫总量的23.33％，31.44％和21.67％；而在柑桔上仅占幼虫总量的2.22％。成虫和幼虫均对受害与未受害果实的选择特性表现出一定的差异，在受害果实上的虫口数量较多，未受害果实上的虫口数量较少。因此，在这6种寄主植物中，南亚果实蝇喜好选择在黄瓜、南瓜和丝瓜以及受害的果实上取食和产卵。
土壤相对水分含量(SRWC)以及空气相对湿度(RH)对南亚果实蝇蛹的生长发育和失水动态均有影响。占78.83％的老熟幼虫集中在SRWC 20-60％的土壤中化蛹。当SRWC为100％时，蛹不能羽化；当SRWC为40％和60％时，蛹具有较短的发育历期(7.54d和7.65d)和较高的羽化率(83.58％和93.3％)。在0％，52％，76％和100％这4个RHs下，蛹的水分损失率随着处理时间的延长而增加，7日蛹水分损失率分别达到63.521％(RH0％)，45.42％(RH52％)，34.23％(RH76％)和10.99％(RH100％)。在这4个RHs下，蛹的发育历期和羽化率亦存在显著差异，在RH100％时，具有较短的发育历期(8.07d)和较高的存活率(90.55％)；而RH0％时，仅3.03％的蛹能够羽化。因此，SRWC 100％和RH0％均不利于南亚果实蝇蛹的发育和存活。
4不同食料对南亚果实蝇生长发育及酶学特性的影响
在室内温度28±0.5℃、相对湿度RH 70±5％、光周期14 L：10 D条件下，研究了南亚果实蝇分别取食黄瓜、南瓜、丝瓜、苦瓜、柑桔和人工饲料时的生长发育特性和繁殖能力。结果表明，南亚果实蝇取食以上各种食料后，幼虫的发育历期和存活率均存在显著差异，分别为4.58d，5.08d，4.50d，6.15d，5.38d，5.72d和86.83％，83.39％，81.19％，51.17％，45.67％，30.33％；成虫的产卵量分别为99.56，84.67，85.91，33.97，8.24和3.87粒，表明取食黄瓜、南瓜和丝瓜后成虫的产卵量较高，取食人工饲料的较低。南亚果实蝇分别取食以上6种食料后，内禀增长率分别为0.0844，0.0572，0.0601，0.0604，-0.0014和-0.0159；种群的趋势指数分别为31.69，20.78，22.62，3.52，0.54和0.11。以上结果均表明，黄瓜、南瓜和丝瓜是该种群生长发育的适宜食料。
超氧化物歧化酶(SOD)、过氧化物酶(POD)和过氧化氢酶(CAT)是生物体内清除活性氧自由基的多酶复合体系，测定了南亚果实蝇分别取食黄瓜、南瓜、冬瓜、丝瓜和苦瓜后，幼虫、蛹和成虫体内SOD、POD和CAT的活性。结果表明，南亚果实蝇幼虫、蛹和成虫体内SOD、POD和CAT的活性均以取食苦瓜时较高，取食冬瓜时次之，取食黄瓜和南瓜时则较低。SOD的活性以幼虫阶段较高，成虫阶段较低；POD和CAT的活性均以成虫阶段较高，蛹阶段较低。表明南亚果实蝇幼虫取食适宜的食料，虫体抗氧化酶的活性较低；而取食相对不适合的食料时，抗氧化酶的活性提高。
羧酸酯酶(CarE)、酸性磷酸酯酶(ACP)、碱性磷酸酯酶(ALP)、细胞色素氧化酶(P450s)和谷胱甘肽S-转移酶(GST)是生物体内降解内源和外源性有毒化合物的重要解毒酶，研究了5种食料对南亚果实蝇这些解毒酶活性的影响。双因子方差分析表明，南亚果实蝇取食黄瓜、南瓜、丝瓜、冬瓜和苦瓜后，解毒酶活性存在显著差异。其中，羧酸酯酶(CarE)活性在黄瓜和南瓜上较高，细胞色素氧化酶P450 O-脱甲基和谷胱甘肽S-转移酶(GST)活性在苦瓜上较高，酸性磷酸酯酶(ACP)和碱性磷酸酯酶(ALP)活性却分别在黄瓜和南瓜上较低。幼虫、蛹和成虫解毒酶的活性亦存在显著差异，成虫具有较高的CarE活性；幼虫具有较高的细胞色素P450 O-脱甲基、GST和ALP活性，但具有较低的ACP活性：除ACP外，蛹的解毒酶活性均较低。因此，南亚果实蝇解毒酶活性受寄主果实种类以及该种群本身发育阶段的影响。
多酚氧化酶是昆虫生命活动过程中的重要调节酶，研究了南亚果实蝇以南瓜为食料时，不同虫态多酚氧化酶的活性和性质。以邻苯二酚为底物，南亚果实蝇不同虫态多酚氧化酶活力大小依次为：3龄幼虫＞成虫＞2龄幼虫＞1龄幼虫＞蛹。在pH6.5，温度34-37℃时的活性较高；在40℃以上时，随着温度的升高酶活性下降。2龄幼虫的K_m和V_(max)较大，表明多酚氧化酶对底物的亲和力较弱，催化能力较强。而以L-多巴为底物时，成虫K_m和3龄幼虫V_(max)较大，表明成虫多酚氧化酶对底物的亲和力较弱，3龄幼虫多酚氧化酶对底物的催化能力较强。因此，南亚果实蝇体内多酚氧化酶的活性和性质与个体的发育阶段密切相关。
5温度对南亚果实蝇生长发育及能源物质含量的影响
以南瓜为食料，研究了6个温度梯度对南亚果实蝇种群生长发育和繁殖的影响。从19℃至28℃，南亚果实蝇幼虫的发育历期随温度的升高而缩短，存活率则随温度的升高而增加；当温度高于28℃时，发育历期随温度的升高而延长，存活率则降低。从19℃至32℃，成虫寿命均随温度的升高而缩短，产卵量在28℃时较高。生殖力生命表参数和种群趋势指数均表明28℃时是南亚果实蝇生长发育和繁殖的适宜温度。
探讨了42℃高温胁迫不同时间对南亚果实蝇成虫寿命以及虫体能源物质即多糖、甘油三酯和可溶性蛋白含量的影响。结果表明，连续处理10d，以每天胁迫10min时成虫寿命较长；连续处理15d时，则以每天胁迫5min时成虫寿命较长。高温胁迫导致虫体多糖贮存量下降，连续处理15d，每天分别胁迫为5，15，30和60min后，多糖含量的损失百分率分别达到为2.04％，2.08％，5.04％和5.93％。对于甘油三酯，当胁迫时间为5min时，虫体甘油三酯的含量较低，此后随着胁迫时间的延长，虫体甘油三酯含量的累积却增加。高温胁迫也会促进南亚果实蝇对蛋白质的利用，随着胁迫时间的延长，虫体蛋白质的贮存量下降。以上结果可以推出，高温胁迫将提高南亚果实蝇虫体对多糖和蛋白质的消耗，而增加甘油三酯的累积。
本文研究了重庆地区南亚果实蝇优势种群的生物生态学特性以及环境因子胁迫后体内多种化学物质含量和活性的变化规律，揭示了南亚果实蝇在重庆地区的发生规律、成灾机理以及适应环境食料和温度的生化机理。研究结果不仅为南亚果实蝇的发生预测和综合治理提供必要的科学依据，亦为南亚果实蝇的分子生物学研究奠定重要的理论基础。
The fruit fly Bactrocera tau Walker, belonging to the order Diptera and the family Tephritidae, mainly infests Cucumis L., Cucurbita L., Luffa L., Benincasa Savi, Momordica L. and other fruits and vegetables. It has taken place in tropic and subtropic regions such as China, Japan, Thailand, Malaysia, Laos, Philippines, Cambodia, India, Bhutan, Indonesia, Sri Lanka and has caused enormous economic losses to fruit and vegetable growers. In China, the fruit fly caused Siraitia grosvenorii Swingle damage up to 21-34% in Guangxi province in 1990 and was injurious to various types of vegetables in Chongqing district in 2004. Due to its economic importance, the study on biological and biochemical characters of this fruit fly was very necessary in the course of the production of fruit and vegetable crops. Based on the research status, the study on the population biological and biochemical characters, especially the physiological mechanisms of the fruit fly adaptable to diets and environmental temperature was carried out systematically. This study was supported by the Specialized Research Fund for the Doctoral Program of Higher Education (20050625005) and the Basic Research Foundation of Chongqing Science Committee (7468), and lasted almost 3 years. The main results are summarized as follows.
1 The species richness and geographical distribution of fruit flies in Chongqing
By the Regional Fruit Fly Project in 2005-2006, systematic trapping using methyl eugenol, cue-lure and hydrolytic protein had been conducted in almost 30 regions of Chongqing. The total 174879 fruit fly samples trapped had confirmed the presence of Bactrocera tau Walker, B. diaphora Hendel, B. scutellata Hendel, B. caudate Fabricius, B. minax Enderlein, B. cucurbitae Coquillett, B. dorsalis Hendel and Dacus trimacula. Among them, B. cucurbitae, B. dorsalis and Dacus trimacula was firstly found in Chongqing district.
Species relative abundance of fruit flies in Chongqing region was analyzed by dominance indexes. The dominance indexes of these fruit flies varied greatly among the 8 kinds of species. In 2005, the dominance indexes of B. tau, B. diaphora, B. scutellata, B. caudate, B. minax, B. cucurbitae and B. dorsalis were 0.8225, 0.1333, 0.0376, 0.0055, 0.0009, 0.0001 and 0.0001, respectively. In 2006, they were 0.7651, 0.0889, 0.1304, 0.0003, 0.0028, 0.0104, 0.0010 and 0.0011 (Dacus trimacula), respectively. The result indicated that B. tau was the dominant population in Chongqing region, B. diaphora and B. scutellata were the second great population, and the others were relatively low in quantities.
According to the identification result of samples collected from different regions of Chongqing, they showed that B. tau, B. diaphora and B. scutellata widely distributed in Chongqing district, but B. minax, B. caudate and D. trimacula were found just in several locations. For B. dorsalis, 9 individuals were captured and distributed in Jiangbei, Yongchuan, Xiushan, Dianjiang and Fuling in 2005, and 95 individuals were captured and distributed in 16 regions including Jiangbei, Jiulongpo, Beibei, and so on in 2006. It showed that the distribution range obviously expanded in 2006. For B. cucurbitae, 10 individuals were trapped in 2005 and 956 individuals were trapped in 2006, and it indicated that the quantity captured in 2006 was significantly higher than that in 2005.
2 The damage character and risk analysis of B. tau population
By the field investigation and the previous study, we comprehensively evaluated the damage character and risk of B. tau. The result showed that B. tau mainly infected cucumber, pumpkin, towel gourd, white gourd, balsam pear and other fruits and vegetables in Chongqing district. Host surveys for this fruit fly in field had confirmed that the damage rate was highest on pumpkin and towel gourd, then on cucumber, and lowest on balsam pear among the five kinds of host fruits above-mentioned. Based on the system and indices of pest risk analysis, we estimated the risk of this fruit fly, and found that the index value of pest risk analysis (R) attained to 1.96. From the result, it could be inferred that this fruit fly was dangerous to some fruits and vegetables, and had posed a threat to other regions in China.
3 The population character of B. tau and its relation with environmental humidity
B. tau had 3-5 generations a year and the generations overlapped in Chongqing district. It was divided into 3 instars in larval stages, and overwintered as pupae in loose soils. Generally, adults mated for several times in the whole life, and the female laid its eggs in clutches under the skin of fruits. The larva or maggot fed on the flesh of fruits. In the process, bacteria were introduced into the fruit causing the fruit to break down or rot. More than 70% mature larvae pupated in the soil with 0-5cm depth, and the survival rate of pupae was higher than 88%. There were 3.74% individuals digging into the soil with 10-25cm depth, and the survival rate was lower than 65%. Only 3.33% individuals could survive in the soil depth over 25cm. The adult could emerge at every time of a day, but mostly aggregated on the time 6: 00-10: 00.
The taxis response and selective propensity of B. tau to six types of fruits cucumber, pumpkin, towel gourd, white gourd, balsam pear and orange were evaluated under indoor experimental conditions. The result demonstrated that most of the adults would obviously prefer to aggregate on cucumber, pumpkin and towel gourd to gain nutrition or lay eggs. Among them, 23.68%, 19.94% and 21.18% of individuals gathered to cucumber, pumpkin and towel gourd, and 21.57%, 18.93% and 22.74% of eggs were lay on the same three host fruits, respectively. It was suitable, adopting linear regress equation, to describe the relationship between the mean quantities and the mean oviposition quantities. As adults, most larvae also aggregated on cucumber (23.33%), pumpkin (31.44%) and towel gourd (21.67%), but little individuals aggregated on orange (2.22%). In addition, the taxis response of the fruit fly to damaged and undamaged fruits was also different in the six kinds of host fruits, and＞50% of individuals would gather to the damaged fruits. From these results, it could seen that the fruit fly preferred to aggregate on cucumber, pumpkin, towel gourd and damaged fruits, but little on orange to gain nutrition or lay eggs.
The effect of soil relative water content (SRWC) and air relative humidity (RH) on the pupal development and survival and the water loss dynamics of B. tau were evaluated under indoor experimental conditions. The pupating selectivity of mature larvae, the pupal duration and eclosion rate varied significantly in the soils with SRWC 0%, 20%, 40%, 60%, 80% and 100%. Most of the mature larvae aggregated on the soils with SRWC 20-60% although they could become pupae in the soils with different water content. When SRWC attained to 100%, all of the pupae failed to survive; SRWC 40% and 60% were most suitable for the pupal development, with the shortest pupal duration (7.54d and 7.65 d) and the highest eclosion rate (83.58% and 93.3%). In the 4 air RHs of 0%, 52%, 76% and 100%, the pupal water loss rates increased with the increase of the treatment time, and reached up to 63.521% for RH 0%, 45.42% for RH 52%, 34.23% for RH 76% and 10.99% for RH 100% at the last day before eclosion. It was suitable, adopting the quadratic equations, to describe the relationship between the water loss rate of pupae accumulated by day and pupal age in day. The pupal duration and eclosion rate were also significantly different in the 4 air RHs. The shortest duration (8.07d) and the highest survival rate (90.55%) of pupae were attained in the air RH 100%, and little individuals could survive in the air RH 0%. The results implied that high soil water content and low air humidity were unfit for the development and survival of B. tau pupae.
4 The influence of diets on the growth and enzyme characterization of B. tau
The effect of different diets including cucumber, pumpkin, towel gourd, balsam pear, orange and artificial diet on the growth and fecundity of B. tau were evaluated at 28±0.5℃, RH 70±5% and photoperiod of 14:10 hours (L: D). The result demonstrated that the developmental duration and survival rate of larvae and the reproduction rate of adults differed significantly among the 6 kinds of diets. When the larva were reared on the above diets, the larval durations were 4.58d, 5.08d, 4.50d, 6.15d, 5.38d, 5.72d, the survival rates were 86.83%, 83.39%, 81.19%, 51.17%, 45.67%, 30.33%, and the total reproduction quantities per female were 99.56, 84.67, 85.91, 33.97 and 8.24, respectively. The result suggested that the larval durations were shorter, the survival rates were higher and the reproduction quantities per female were also higher on cucumber, pumpkin and towel gourd. According to the life-table parameters and the population trend indexes, it could be concluded that cucumber, pumpkin and towel gourd were the most suitable diets for the development and fecundity of this fruit fly, but contrary for orange and artificial diet.
Superoxide dismutase (SOD), Peroxidase (POD) and Catalase (CAT) are considered to be an important physiological and biochemical mechanism of organisms against adverse environmental factors. The soluble protein contents and those enzyme activities in B. tau fed on cucumber, pumpkin, towel gourd, white gourd and balsam pear were determined by biochemical methods. The result indicated that the protein contents and those detoxification enzyme activities varied significantly when the fruit fly were reared on different diets—cucumber, pumpkin, towel gourd, white gourd and balsam pear. Among the 5 kinds of diets, the protein contents in this fruit fly were highest on balsam pear while lowest on cucumber and white gourd. The SOD, POD and CAT activities were highest on balsam pear, then white gourd, but lowest on cucumber and pumpkin, and those enzyme activities also varied greatly in different developmental stages of this fruit fly. The SOD activity was highest at the larval stage, and lowest at the adult stage; the POD and CAT activities were highest at the adult stage, but lowest at pupal stage. Based on the above study, it indicated that the antioxidant activity in B. tau was lower when the fruit fly was reared on suitable diets, than that on unsuitable diets.
This study was conducted to determine whether the activity of detoxification enzymes, such as carboxylesterase (CarE), acid phosphatase (ACP), alkaline phosphatase (ALP), cytochrome P450-dependent O-demethylase and glutathione S-transferase (GST) in Bactrocera tau (Walker), respond to different diets and developmental stages of this fruit fly. Two-way ANOVA showed that protein contents and these detoxification enzyme activities varied significantly when the fruit fly were reared on different diets cucumber, pumpkin, towel gourd, white gourd and balsam pear. Among the five kinds of diets, protein contents in the fruit fly were highest on balsam pear while lowest on cucumber and white gourd. The CarE activity was highest on cucumber and pumpkin, cytochrome P450-dependent O-demethylase and GST activities were highest on balsam pear, but ACP and ALP activities were lowest on cucumber and pumpkin, respectively. In addition, these detoxification enzyme activities were also significantly different in the larval, pupal and adult stages of the fruit fly. Adults had the highest CarE activity, and larvae had the highest ALP, cytochrome P450-dependent O-demethylase and GST activities, but the lowest ACP activity. Generally, these detoxification enzyme activities in the pupal stage were lower than those in the larval and adult stages except for ACP. The result indicated that these detoxification enzyme activities of B. tau were closely related to host plant species and developmental stages of this fruit fly.
Polyphenoloxidase (PPO), a group of copper proteins thatare widely distributed from bacteria to mammals, catalyzes the oxidation of hydroxyphenols to their quinone derivatives, which then spontaneously polymerize. The kinetic properties of PPO from B. tau different stages were compared and determined by a series of chemical methods. The result showed that PPO activity differed significantly at different stages in this insect. The PPO activity of the 3rd instar was highest and that of pupae was lowest among the 1st instar, the 2nd instar, the 3rd instar, pupae and adults. The order of the enzyme activity was the 3 instar＞adults＞the 2nd instar＞the 1st instar＞pupae. The optimum pH was 7.0 and the best temperature was 34-37℃for the tested PPO. The kinetic parameter for the oxidation of catechol by PPO from the different stages was determined and compared, and it showed that the affinity of PPO to the substrates catechol was higher, but the catalytic activity was lower for the 2nd instar. At the same time, for the substrates L-DOPA, the K_m for adults and V_(max) for the 3rd instar of PPO were highest, it showed that the affinity of PPO for adults was lower, and the catalytic activity for the 3rd instar was higher than the other stages. From the above conclusion, it could be inferred that the property of PPO activities is closely associated with the developmental stages of B. tau.
5 The influence of temperature on the growth and energy resources of B. tau
The growth and fecundity of B. tau fed on pumpkin were investigated at 6 constant temperatures 19℃, 22℃, 25℃, 28℃, 31℃and 34℃in a series of indoor trials. ANOVA showed that temperature affected significantly the developmental duration and reproduction capacity. The developmental duration of larvae varied from 6.31d at 28℃to 10.09d at 19℃, and the survival rate varied from 27.03% at 34℃to 74.76% at 28℃. The female adult at 19℃and 22℃had significantly greater longevity (96.29d and 94.00d) than those at the other temperature, and had the greatest reproduction capacity (105.91 eggs per female) at 28℃. Accordingly, the intrinsic rate of increase (r_m) and the population trend index were highest (0.0572 and 17.28, respectively) for this fruit fly at 28℃. The results implied that the temperature of 28℃were the most suitable for the population development and fecundity.
The longevity and energy sources including polysaccharides, triacylglycerol and soluble protein of B. tau submitted to heat shocks (42℃) of various durations (5,15, 30 or 60 min daily) for 10d or 15d were determined systematically by ecological and biochemical methods. A slight longevity increase was observed in 15min daily for 10d and in 5min daily for 15d, but longer shocks had negative effects. Fruit flies submitted to the procedure providing a longevity increase did not show a higher polysaccharide and protein contents than those in the other treatment durations. However, the highest polysaccharide and protein contents were observed in the control flies. Except for the control flies, the accumulation of triacylglycerol of this fruit fly increased responding to the treated durations. The above results suggested that under the heat shocks conditions, polysaccharides and soluble protein were the main energy sources, while the utilization of triacylglycerol decreased.
Generally, the study on the bio-ecological characteristics, and the influence of diets and heat shock on the physiological and biochemical properties of B. tau has clearly elucidated the occurrence regulation, formative mechanism and biochemical mechanism of this fruit fly suitable to diets and temperature. The result will provide necessary information for IPM program, and enrich the whole theoretical system of fruit flies.
引文
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