家蝇与大头金蝇幼虫取食与成虫产卵行为研究
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摘要
随着近年国际上鱼粉产量的减少和鱼粉价格的上涨,人们不断寻求能够代替鱼粉的植物或动物源蛋白。营养价值很高的蝇蛆已被证实能够替代鱼粉用于动物养殖业。但是,较高的饲料生产成本阻碍了蝇蛆的大规模养殖。因此,详尽地研究蝇类的食性和产卵选择行为,改善蝇蛆饲料的理化特性,对于蝇类的经济饲养具有重要的意义。本论文对家蝇和大头金蝇幼虫的取食和选择行为,以及成虫的产卵选择行为进行了研究。具体结论如下:
使用瘦肉、麦麸和鱼粉饲养大头金蝇和家蝇初孵化的幼虫时,除了大头金蝇幼虫不能在麦麸上存活外,两种幼虫在瘦肉、鱼粉,以及家蝇在麦麸上都能正常生长发育至成虫。与麦麸和鱼粉相比,瘦肉饲养的大头金蝇和家蝇幼虫期最短、成虫羽化率最高以及成虫翅最长(p<0.01)。部分2龄或3龄大头金蝇能在麦麸上独立发育至成虫。但是当与2龄或3龄家蝇幼虫在麦麸上混合饲养时,大头金蝇的生长状况受到抑制,不能正常化蛹。2龄家蝇幼虫对麦麸和瘦肉的选择性差异不显著,2龄大头金蝇幼虫对瘦肉的选择性显著高于麦麸(p<0.01)。与新鲜食物相比,家蝇或大头金蝇幼虫都明显选择取食幼虫取食过的食物(p<0.01)。在3种基质中,家蝇雌虫最嗜在麦麸中产卵,而大头金蝇最嗜在瘦肉中产卵(p<0.01)。
食物的蛋白含量和幼虫的种群密度能影响野外家蝇和大头金蝇种群F_1代幼虫的种间竞争和种内竞争的强度。当食物的蛋白含量不小于4.6%时,大头金蝇和家蝇幼虫的种间竞争作用大于家蝇幼虫种内作用,而小于大头金蝇幼虫的种内竞争作用。当家蝇和大头金蝇幼虫的饲养密度升高时,蝇的发育历期延长、存活率降低、成虫体重降低或翅长缩短(p<0.05)。不同于实验室种群,使用麦麸饲养野外家蝇和大头金蝇F_1代幼虫混合种群时,与用麦麸单独饲养或用水饲养的同龄大头金蝇幼虫相比,用麦麸饲养的混合种群中能正常羽化至成虫的大头金蝇发育历期缩短、存活率升高、成虫体重或翅长增加(p<0.05),而与麦麸单独饲养的家蝇相比,混合种群中的家蝇发育历期缩短、存活率降低、体重降低或翅长减小。
当用麦麸作为营养源时,麦麸的含水量对野外家蝇和大头金蝇幼虫的生长发育影响显著。家蝇初孵幼虫能在55%～75%湿麦麸上正常生长,2龄大头金蝇幼虫能在45%～75%的湿麦麸上发育至成虫。在食物充足的情况下,麦麸的湿度对家蝇的发育历期没有显著影响,而对存活率、雌成虫体重和翅长有显著影响(p<0.01)。麦麸湿度对大头金蝇的的发育历期,存活率、雌成虫体重和翅长都有显著影响(p<0.01)。
野外家蝇和大头金蝇F_1代成虫在麦麸、瘦肉以及两者混合物中产卵的量存在显著差异(p<0.01)。无论幼虫用麦麸或瘦肉饲养,家蝇都显著选择在麦麸和瘦肉的混合物产卵,仅在麦麸上产少量的卵,不在瘦肉上产卵。家蝇幼虫期的取食经历对成虫的产卵基质选择行为没有影响。幼虫用瘦肉饲养的大头金蝇明显选择在瘦肉以及麦麸和瘦肉的混合物上产卵,两者之间差异不显著,不在麦麸上产卵。
麦麸的含水量和粒度对家蝇产卵基质选择行为影响显著(p<0.01)。产卵家蝇分别在粒度大于2mm或含水量为60%的湿麦麸中产卵最多。在嗅觉仪实验中,以水为对照时,麦麸或瘦肉对家蝇和大头金蝇雌、雄蝇表现出较强的引诱性(p<0.01)。在对麦麸和瘦肉双向选择时,家蝇和大头金蝇的雌、雄蝇都显著选择向瘦肉定位飞行(p<0.01),另外雌、雄大头金蝇在瘦肉中分布的数量存在显著差异,雌蝇对瘦肉的定位更加显著(p<0.05),而家蝇两性之间的差异不显著。在麦麸中添加各种氨基酸不能增加麦麸对大头金蝇的产卵引诱力,大头金蝇雌蝇仍将全部的卵产在瘦肉中。家蝇在麦麸中的产卵量要显著地高于在瘦肉以及葡萄糖、蔗糖或淀粉处理的瘦肉的产卵量,各种处理的瘦肉中的卵量差异不显著(p<0.05)。
同种幼虫的存在不能提高雌家蝇在湿麦麸中的产卵量,而同种幼虫取食过的产卵基质对雌家蝇有很强的产卵引诱作用。除了鱼粉饲养的大头金蝇将全部的卵产在幼虫取食过的鱼粉上外,家蝇和大头金蝇在幼虫取食过的基质上的产卵数量显著多于在新鲜基质上的产卵量。在整个产卵期内,日龄对大头金蝇和家蝇的产卵基质选择行为没有明显影响。大头金蝇只选择在瘦肉上产卵,而家蝇明显偏嗜在麦麸中产卵(p<0.01),在鱼粉和瘦肉中产卵较少。
取食经历对2龄家蝇幼虫的食物选择行为有显著影响(p<0.01)。在双向选择实验中,幼虫前期用麦麸饲养的2龄家蝇幼虫在麦麸或瘦肉中分布的数量没有显著差异,而具有瘦肉经历的幼虫明显选择取食瘦肉。取食经历并不能改变2龄大头金蝇幼虫的取食选择行为,无论取食鱼粉或瘦肉,幼虫都显著选择取食鱼粉。
不管幼虫或蛹期经历何种产卵基质,羽化后的大头金蝇雌蝇都将全部的卵产在瘦肉上。但是羽化前的经历对家蝇产卵选择行为有显著影响(p<0.01)。当幼虫用麦麸饲养时,与蛹期经历瘦肉或滤纸联合水洗的雌蝇相比,蛹期经历麦麸和滤纸的雌蝇明显地将更多的卵产在麦麸上(p<0.05)。当幼虫用瘦肉饲养时,与蛹期经历滤纸或滤纸联合水洗的雌蝇相比,蛹期经历麦麸的雌蝇明显地将更多的卵产在麦麸上(p<0.05)。幼虫的取食经历对雌家蝇的产卵基质选择行为没有显著影响,而蛹期的基质经历对雌家蝇的产卵基质选择行为有显著的影响(p<0.01)。与蛹期经历瘦肉、滤纸或者滤纸联合水洗的雌家蝇相比,蛹期经历麦麸的雌家蝇明显地将更多的卵产在麦麸上(p<0.01)。
家蝇雌蝇产卵器和足跗节的超微结构观察显示,在肛上板、肛下板、肛尾叶和跗节上存在数量不等的感受鬃、刺形感受器、毛形感受器、锥形感受器、腔锥形感受器和腔形感受器。刺形感受器在跗节上分布广泛,毛形感受器在产卵器和跗节上分布广泛。跗节、前跗节和产卵器上的感受鬃、刺形感受器和毛形感受器具有机械感受器典型的形态特征,有助于雌蝇感受产卵基质的疏松度。肛尾叶末端的栓锥形感受器可能与湿度感受有关。
Scarcity and cost of fish meal as a livestock and commercial fish food has increased the need to find alternative plant and animal protein based foods.Fly larvae are used to produce Magmeal.This food supplement has high nutritional value and has successfully replaced fishmeal in the diet of animals.However,popularization of the magmal in the poulry and fishery is faced with problem of the high cost of the diet of the fly larvae.To improve the chemical and physical characters of the fly diet,in this paper,we designed several tests to reveal the mechanisms of the larval feeding and adult oviposition
preference of Musca domestica L.(MD) and Chrysomya megacephala(F.)(CM). In the first test,larval food preferences,development,and oviposition preference of 10-15 days old MD and CM were evaluated on three substrates:wheat bran,pork muscle, and fish meal.Pork muscle was the optimal food for both species.MD larvae completed development on the wheat bran,but CM did not.In choice tests,the second instar CM larvae preferred pork muscle to wheat bran(p<0.05),while second instar MD larvae showed no preference.Female CM predominantly chose pork muscle as-the oviposition substrate.Female MD laid eggs on three substrates and wheat bran showed the greatest attrativeness to gravid MD(p<0.01).
The second test is to evaluate the larval nutrition resource competition between the F_1 larvae of CM and MD wild populations.The results indicated that the extreme nutrition restriction produced shorter MDor CM pre-adult development time,but MD or CM exhibited longer pre-adult development time and lower body weight and smaller size of following adults when larvae were reared on nutrition substrates with varying protein contents(p<0.05).CM larvae exhibited stronger resource competition potential than MD larvae on nutrition materials with the content of protein >6.4%.The development of second or third instar CM larvae on wheat bran could be improved significantly when reared with second or third instar MD larvae,which would help CM to extend their ecological niches into plant materials.The second instar CM larvae could enter into the pupal state on the wet bran with the water content of 45-75%,but the first instar MD only grew normally on 55-75%water content of wet wheat bran.The F_1 females of MD and CM all preferred to lay their eggs on the mixtures of the wheat bran and pork muscle than wheat bran or pork muscle.With the increasing of animal protein substrates in garbage in the urban-suburban areas,CM has the potential of displacing the established house fly from this ecological niche in some urban-suburban areas.
The third test was to evlaute the effect of the chemical and physical character of the oviposition substrate,the age of the female and existence of the conspecific larvae on the oviposition preference of female and/or MD and/or CM.Female MD significantly preferred ovipositing on wet wheat bran with the water content of 60%or with a particle size of > 2.0 mm dia(p<0.01).In the two-choice olfactometer,wheat bran or pork muscle was more attractive to female and male MD or CM than distilled water(p<0.01),and pork muscle showed greater attractiveness to female and male MD or CM than wheat bran (p<0.01).Female CM was more attracted by pork muscle than male CM(p<0.05),which was not found in MD(p>0.05).The acceptability of wheat bran to gravid CM was not enhanced with the addition of different acid aminos and MD females did not lay more eggs on pork mucle when glucose,sucrose and starch were added.MD females did not lay more eggs on the wheat bran with the existence of conspecific larvae.But the food which had been ingested by conspecific larvae showed greater attractiveness to female MD or CM than fresh food.During the whole oviposition period,age had no significant effect on the oviposition preference of MD and CM.MD females laid more eggs on wheat bran than on fishmeal and pork muscle(p<0.01),and CM females always laid their eggs on pork muscle.
The fourth test are to evaluate the effects of larval experience on larval feeding preference and the effects of larval and pupal experience on oviposition preference of females in MD and CM.Dietary experience since hatch to test could significantly influence feeding preference of the second instar larvae in MD(p<0.01),but could not in CM.The larval dietary and pupal experience did not change the oviposition preference of CM and experienced gravid females laid all of their eggs on pork muscle.Larval feeding regimen had no effect on oviposition preference of subsequent females in MD.However, the pupal experience in MD significantly influenced the oviposition preference of subsequent females,and MD females from pupae matured on wheat bran laid significantly more eggs on wheat bran than that matured on pork muscle or filter paper whether pupae were washed or not(p<0.01).These results demonstrated that host-selection behavior in MD was shaped by preimaginal experience according to the chemical legacy hypothesis.
In the last test,the fine structure of the ovipositor and tarsi of front,middle and hind legs of female MD were studied using scanning electron microscopy to contribute detailed morphological information on these organs of this species.Several types of sensilla were found on the tarsi,hypoproct,epiproct and cercus,including the sensilla bristle,sensilla chaetica,sensilla trichodea,sensilla basiconica,sensilla coeloconica and sensilla cavity. The bristle,sensilla chaetica and sensilla trichodea with typical morphological characters of mechanoreceptors may provide mechanical information of the oviposition substrate. The sensilla styloconica on the terminal of the dorsal side of the left cercus may serve as hydroreceptor of females.
使用瘦肉、麦麸和鱼粉饲养大头金蝇和家蝇初孵化的幼虫时,除了大头金蝇幼虫不能在麦麸上存活外,两种幼虫在瘦肉、鱼粉,以及家蝇在麦麸上都能正常生长发育至成虫。与麦麸和鱼粉相比,瘦肉饲养的大头金蝇和家蝇幼虫期最短、成虫羽化率最高以及成虫翅最长(p<0.01)。部分2龄或3龄大头金蝇能在麦麸上独立发育至成虫。但是当与2龄或3龄家蝇幼虫在麦麸上混合饲养时,大头金蝇的生长状况受到抑制,不能正常化蛹。2龄家蝇幼虫对麦麸和瘦肉的选择性差异不显著,2龄大头金蝇幼虫对瘦肉的选择性显著高于麦麸(p<0.01)。与新鲜食物相比,家蝇或大头金蝇幼虫都明显选择取食幼虫取食过的食物(p<0.01)。在3种基质中,家蝇雌虫最嗜在麦麸中产卵,而大头金蝇最嗜在瘦肉中产卵(p<0.01)。
食物的蛋白含量和幼虫的种群密度能影响野外家蝇和大头金蝇种群F_1代幼虫的种间竞争和种内竞争的强度。当食物的蛋白含量不小于4.6%时,大头金蝇和家蝇幼虫的种间竞争作用大于家蝇幼虫种内作用,而小于大头金蝇幼虫的种内竞争作用。当家蝇和大头金蝇幼虫的饲养密度升高时,蝇的发育历期延长、存活率降低、成虫体重降低或翅长缩短(p<0.05)。不同于实验室种群,使用麦麸饲养野外家蝇和大头金蝇F_1代幼虫混合种群时,与用麦麸单独饲养或用水饲养的同龄大头金蝇幼虫相比,用麦麸饲养的混合种群中能正常羽化至成虫的大头金蝇发育历期缩短、存活率升高、成虫体重或翅长增加(p<0.05),而与麦麸单独饲养的家蝇相比,混合种群中的家蝇发育历期缩短、存活率降低、体重降低或翅长减小。
当用麦麸作为营养源时,麦麸的含水量对野外家蝇和大头金蝇幼虫的生长发育影响显著。家蝇初孵幼虫能在55%～75%湿麦麸上正常生长,2龄大头金蝇幼虫能在45%～75%的湿麦麸上发育至成虫。在食物充足的情况下,麦麸的湿度对家蝇的发育历期没有显著影响,而对存活率、雌成虫体重和翅长有显著影响(p<0.01)。麦麸湿度对大头金蝇的的发育历期,存活率、雌成虫体重和翅长都有显著影响(p<0.01)。
野外家蝇和大头金蝇F_1代成虫在麦麸、瘦肉以及两者混合物中产卵的量存在显著差异(p<0.01)。无论幼虫用麦麸或瘦肉饲养,家蝇都显著选择在麦麸和瘦肉的混合物产卵,仅在麦麸上产少量的卵,不在瘦肉上产卵。家蝇幼虫期的取食经历对成虫的产卵基质选择行为没有影响。幼虫用瘦肉饲养的大头金蝇明显选择在瘦肉以及麦麸和瘦肉的混合物上产卵,两者之间差异不显著,不在麦麸上产卵。
麦麸的含水量和粒度对家蝇产卵基质选择行为影响显著(p<0.01)。产卵家蝇分别在粒度大于2mm或含水量为60%的湿麦麸中产卵最多。在嗅觉仪实验中,以水为对照时,麦麸或瘦肉对家蝇和大头金蝇雌、雄蝇表现出较强的引诱性(p<0.01)。在对麦麸和瘦肉双向选择时,家蝇和大头金蝇的雌、雄蝇都显著选择向瘦肉定位飞行(p<0.01),另外雌、雄大头金蝇在瘦肉中分布的数量存在显著差异,雌蝇对瘦肉的定位更加显著(p<0.05),而家蝇两性之间的差异不显著。在麦麸中添加各种氨基酸不能增加麦麸对大头金蝇的产卵引诱力,大头金蝇雌蝇仍将全部的卵产在瘦肉中。家蝇在麦麸中的产卵量要显著地高于在瘦肉以及葡萄糖、蔗糖或淀粉处理的瘦肉的产卵量,各种处理的瘦肉中的卵量差异不显著(p<0.05)。
同种幼虫的存在不能提高雌家蝇在湿麦麸中的产卵量,而同种幼虫取食过的产卵基质对雌家蝇有很强的产卵引诱作用。除了鱼粉饲养的大头金蝇将全部的卵产在幼虫取食过的鱼粉上外,家蝇和大头金蝇在幼虫取食过的基质上的产卵数量显著多于在新鲜基质上的产卵量。在整个产卵期内,日龄对大头金蝇和家蝇的产卵基质选择行为没有明显影响。大头金蝇只选择在瘦肉上产卵,而家蝇明显偏嗜在麦麸中产卵(p<0.01),在鱼粉和瘦肉中产卵较少。
取食经历对2龄家蝇幼虫的食物选择行为有显著影响(p<0.01)。在双向选择实验中,幼虫前期用麦麸饲养的2龄家蝇幼虫在麦麸或瘦肉中分布的数量没有显著差异,而具有瘦肉经历的幼虫明显选择取食瘦肉。取食经历并不能改变2龄大头金蝇幼虫的取食选择行为,无论取食鱼粉或瘦肉,幼虫都显著选择取食鱼粉。
不管幼虫或蛹期经历何种产卵基质,羽化后的大头金蝇雌蝇都将全部的卵产在瘦肉上。但是羽化前的经历对家蝇产卵选择行为有显著影响(p<0.01)。当幼虫用麦麸饲养时,与蛹期经历瘦肉或滤纸联合水洗的雌蝇相比,蛹期经历麦麸和滤纸的雌蝇明显地将更多的卵产在麦麸上(p<0.05)。当幼虫用瘦肉饲养时,与蛹期经历滤纸或滤纸联合水洗的雌蝇相比,蛹期经历麦麸的雌蝇明显地将更多的卵产在麦麸上(p<0.05)。幼虫的取食经历对雌家蝇的产卵基质选择行为没有显著影响,而蛹期的基质经历对雌家蝇的产卵基质选择行为有显著的影响(p<0.01)。与蛹期经历瘦肉、滤纸或者滤纸联合水洗的雌家蝇相比,蛹期经历麦麸的雌家蝇明显地将更多的卵产在麦麸上(p<0.01)。
家蝇雌蝇产卵器和足跗节的超微结构观察显示,在肛上板、肛下板、肛尾叶和跗节上存在数量不等的感受鬃、刺形感受器、毛形感受器、锥形感受器、腔锥形感受器和腔形感受器。刺形感受器在跗节上分布广泛,毛形感受器在产卵器和跗节上分布广泛。跗节、前跗节和产卵器上的感受鬃、刺形感受器和毛形感受器具有机械感受器典型的形态特征,有助于雌蝇感受产卵基质的疏松度。肛尾叶末端的栓锥形感受器可能与湿度感受有关。
Scarcity and cost of fish meal as a livestock and commercial fish food has increased the need to find alternative plant and animal protein based foods.Fly larvae are used to produce Magmeal.This food supplement has high nutritional value and has successfully replaced fishmeal in the diet of animals.However,popularization of the magmal in the poulry and fishery is faced with problem of the high cost of the diet of the fly larvae.To improve the chemical and physical characters of the fly diet,in this paper,we designed several tests to reveal the mechanisms of the larval feeding and adult oviposition
preference of Musca domestica L.(MD) and Chrysomya megacephala(F.)(CM). In the first test,larval food preferences,development,and oviposition preference of 10-15 days old MD and CM were evaluated on three substrates:wheat bran,pork muscle, and fish meal.Pork muscle was the optimal food for both species.MD larvae completed development on the wheat bran,but CM did not.In choice tests,the second instar CM larvae preferred pork muscle to wheat bran(p<0.05),while second instar MD larvae showed no preference.Female CM predominantly chose pork muscle as-the oviposition substrate.Female MD laid eggs on three substrates and wheat bran showed the greatest attrativeness to gravid MD(p<0.01).
The second test is to evaluate the larval nutrition resource competition between the F_1 larvae of CM and MD wild populations.The results indicated that the extreme nutrition restriction produced shorter MDor CM pre-adult development time,but MD or CM exhibited longer pre-adult development time and lower body weight and smaller size of following adults when larvae were reared on nutrition substrates with varying protein contents(p<0.05).CM larvae exhibited stronger resource competition potential than MD larvae on nutrition materials with the content of protein >6.4%.The development of second or third instar CM larvae on wheat bran could be improved significantly when reared with second or third instar MD larvae,which would help CM to extend their ecological niches into plant materials.The second instar CM larvae could enter into the pupal state on the wet bran with the water content of 45-75%,but the first instar MD only grew normally on 55-75%water content of wet wheat bran.The F_1 females of MD and CM all preferred to lay their eggs on the mixtures of the wheat bran and pork muscle than wheat bran or pork muscle.With the increasing of animal protein substrates in garbage in the urban-suburban areas,CM has the potential of displacing the established house fly from this ecological niche in some urban-suburban areas.
The third test was to evlaute the effect of the chemical and physical character of the oviposition substrate,the age of the female and existence of the conspecific larvae on the oviposition preference of female and/or MD and/or CM.Female MD significantly preferred ovipositing on wet wheat bran with the water content of 60%or with a particle size of > 2.0 mm dia(p<0.01).In the two-choice olfactometer,wheat bran or pork muscle was more attractive to female and male MD or CM than distilled water(p<0.01),and pork muscle showed greater attractiveness to female and male MD or CM than wheat bran (p<0.01).Female CM was more attracted by pork muscle than male CM(p<0.05),which was not found in MD(p>0.05).The acceptability of wheat bran to gravid CM was not enhanced with the addition of different acid aminos and MD females did not lay more eggs on pork mucle when glucose,sucrose and starch were added.MD females did not lay more eggs on the wheat bran with the existence of conspecific larvae.But the food which had been ingested by conspecific larvae showed greater attractiveness to female MD or CM than fresh food.During the whole oviposition period,age had no significant effect on the oviposition preference of MD and CM.MD females laid more eggs on wheat bran than on fishmeal and pork muscle(p<0.01),and CM females always laid their eggs on pork muscle.
The fourth test are to evaluate the effects of larval experience on larval feeding preference and the effects of larval and pupal experience on oviposition preference of females in MD and CM.Dietary experience since hatch to test could significantly influence feeding preference of the second instar larvae in MD(p<0.01),but could not in CM.The larval dietary and pupal experience did not change the oviposition preference of CM and experienced gravid females laid all of their eggs on pork muscle.Larval feeding regimen had no effect on oviposition preference of subsequent females in MD.However, the pupal experience in MD significantly influenced the oviposition preference of subsequent females,and MD females from pupae matured on wheat bran laid significantly more eggs on wheat bran than that matured on pork muscle or filter paper whether pupae were washed or not(p<0.01).These results demonstrated that host-selection behavior in MD was shaped by preimaginal experience according to the chemical legacy hypothesis.
In the last test,the fine structure of the ovipositor and tarsi of front,middle and hind legs of female MD were studied using scanning electron microscopy to contribute detailed morphological information on these organs of this species.Several types of sensilla were found on the tarsi,hypoproct,epiproct and cercus,including the sensilla bristle,sensilla chaetica,sensilla trichodea,sensilla basiconica,sensilla coeloconica and sensilla cavity. The bristle,sensilla chaetica and sensilla trichodea with typical morphological characters of mechanoreceptors may provide mechanical information of the oviposition substrate. The sensilla styloconica on the terminal of the dorsal side of the left cercus may serve as hydroreceptor of females.
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