摘要
蝴蝶色彩斑斓具有很高的观赏价值和经济价值，蝴蝶帮助植物授粉，还有重要的生态价值。本文研究了访花蝴蝶和食腐蝴蝶觅食过程中对花色、花形和气味等视觉和嗅觉信号的利用权重，分析了其生理原因，有助于认识访花蝴蝶与显花植物间的生态关系以及食腐蝴蝶取食习性与环境的进化适应关系。主要研究结果如下：
（1）访花蝴蝶觅食过程中对蜜源植物的视觉和嗅觉行为反应
通过对七种访花蝴蝶在觅食过程中对花冠颜色及大小的行为反应观察，从红、橙、黄、紫和白色五种颜色花单独的吸引作用及以其与蜂蜜水结合模式下的引诱作用实验中，我们发现嗅觉和视觉信号对不同蝶种的行为影响显著不同。本实验涉及的七种蝴蝶，除虎斑蝶外，对颜色都有一定的趋性。白带锯蛱蝶和金斑蝶以视觉为主定位寻找食物；红锯蛱蝶、青斑蝶和大帛斑蝶觅食时以嗅觉为主，视觉为辅；迁粉蝶对视觉和嗅觉利用程度相等。我们首次发现，专性访花的虎斑蝶对供试颜色不敏感，靠嗅觉信号觅食。七种蝴蝶觅食过程中对嗅觉和视觉信号利用有较大的区别，有4种途径：1）嗅觉信号为主，视觉信号为辅；2）视觉信号为主，嗅觉信号为辅；3）视觉信号和嗅觉信号同等重要；4）只依靠嗅觉信号。
蝴蝶的觅食过程中，嗅觉信号可能是蝴蝶觅食的关键因子，可能对所有蝴蝶觅食产生引诱作用，视觉可能是蝴蝶发现食物源的重要因子，而后主要靠嗅觉信号来刺激取食。
（2）访花蝴蝶对花的大小的选择
我们所实验的七种蝴蝶，除青斑蝶外，均倾向于花形较大的花朵，可能是大型花朵视觉刺激更为强烈，挥发性物质更多的缘故，也可能与蝴蝶本身的体型有关。七种蝴蝶的体型属于大中型蝴蝶，因此它们对大中型花朵的偏爱可能是出于自身能量的需求。说明蝴蝶的访花策略不止取决于视觉和嗅觉的利用权重，还与其自身体型有关。
（3）食腐蝴蝶觅食过程中对视觉和嗅觉行为反应
我们观察了枯叶蛱蝶成虫对红、黄、紫和白色四种颜色和六种腐烂水果（梨、苹果、香蕉、西瓜、橙子和柿子）汁液的觅食反应。结果发现，在无食物气味情况下，成虫对四种颜色均无觅食反应，当花上喷有腐烂梨汁后，强烈吸引枯叶蛱蝶访食，说明食腐蝴蝶觅食过程中无视觉反应，主要靠嗅觉发现食物。枯叶蛱蝶对六种腐烂水果汁液的偏好无显著差异。
（4）蝴蝶对食物挥发物的田间行为反应及电生理测定
1）青斑蝶对蜜源植物挥发物的田间行为反应
我们在田间观察了青斑蝶对蜜源植物马缨丹和马利筋的花部挥发物α-蒎烯、水杨酸甲酯，花部挥发物类似物1-辛醛，桉叶油醇，萜品醇，以及前人报道的花中普遍存在的苯甲醛的觅食反应。结果发现，无论是六种单一化合物、α-蒎烯、水杨酸甲酯、1-辛醛和桉叶油醇的化合物组合还是α-蒎烯和水杨酸甲酯的不同剂量，青斑蝶对蜜源植物挥发物的取食均未显著高于去离子水，且不同化合物之间无明显差异。
2）枯叶蛱蝶对腐烂水果挥发物的田间行为反应及电生理测定
我们在田间观察了枯叶蛱蝶对腐烂水果挥发物异戊醇、乙酸乙酯、乙酸异戊酯、α-蒎烯，2-戊酮的同系物丁酮和在其他发酵物中常见的两种挥发物乙醇和乙酸的取食反应并测定了新羽化成虫对这些挥发物触角电位反应。田间行为测定发现，醇类对枯叶蛱蝶的引诱最为强烈，其次是酯类。当乙酸乙酯与乙醇混合后，枯叶蛱蝶取食次数增加。枯叶蛱蝶对乙酸乙酯和乙醇两种化合物的访食次数均随浓度的增加而增强。在触角电位反应中，无论是浓度5μl/ml还是50μl/ml，同一性别成虫对异戊醇、乙酸异戊酯、乙酸乙酯、α-蒎烯、丁酮和乙酸等六种不同化合物的相对EAG反应值差异均不显著，且相对EAG值均高于乙醇（100%）。成虫对异戊醇和乙酸在特定浓度下的EAG反应表现出性别差异。当浓度为50μl/ml时，成虫对四种化合物组合的相对EAG值均未表现出性别差异。相同性别成虫在特定浓度范围内（0.01、0.1、1、5、10和100μl/ml）对乙醇或乙酸乙酯的相对EAG值均与挥发物浓度无显著相关性。结果证明，嗅觉信号在枯叶蛱蝶觅食中发挥主要作用，觅食成虫可利用广泛的食物信息化学物质，但成虫对挥发物的取食偏好不一定与触角电位反应相关。
（5）蝴蝶触角感器与嗅觉行为反应分析
应用扫描电镜对访花的虎斑蝶、金斑蝶、大帛斑蝶和迁粉蝶以及食腐的枯叶蛱蝶雌雄成虫触角及其感器外部形态进行了观察和研究。结果表明，五种蝴蝶触角感器种类和形态相同，无明显性二型现象。触角共有五类感器，即B hm氏鬃毛、鳞形感器、毛形感器（Ⅰ、Ⅱ）、腔锥形感器和刺形感器，并对各类型感器的分布和数量进行了描述。五种蝴蝶的感器总数均是雄虫多于雌虫，这可能与雄虫求偶有关。结合五种蝴蝶觅食时的嗅觉利用，我们推测毛形感器Ⅱ和腔锥形感器具有感受食物气味的功能，鳞形感器具辅助作用。具有感受食物气味或辅助感受食物气味的感器数量可以反映蝴蝶嗅觉利用权重。
（6）蝴蝶复眼与觅食行为初步分析
利用光学显微镜和透射电镜对访花的虎斑蝶、金斑蝶、大帛斑蝶和迁粉蝶以及食腐的枯叶蛱蝶雌、雄成虫复眼的小眼数量和复眼内部的显微和超显微结构进行观察。结果表明，五种蝴蝶的雌、雄成虫均具有大量小眼，同一蝶种的小眼数目均是雄虫多于雌虫，但仅迁粉蝶的小眼数量有显著性别差异。在复眼结构上，五种蝴蝶均无性二型差异。结合五种蝴蝶觅食时的视觉利用，我们推测小眼数目基本能反映蝴蝶觅食时的信号利用规律。
Butterflies have important ornamental value and economic value because of their colorfulwings. Meanwhile, butterflies are also playing ecological value during visiting flowers. In thispaper, we found that the relative strength of butterfly response to visual versus olfactory cuesdiffered among butterfly species by observing the foraging responses of flower-feeding andfruit-feeding butterflies and uncovered the related physiology reasons. Understanding theflower-visiting characteristics of butterflies can reveal the ecological relationships betweenbutterflies and flowers. And knowing the fruit-feeding habits of butterflies can reveal theadaption relationships between butterflies and environment. The main conclusions are asfollows:
(1)The visual and olfactory responses during nectar-feeding butterflies foraging
Foraging responses of seven butterfly species were determined for flower models thatdiffered in color, size, and depth of artificial corolla. We measured frequency of visits toflowers of various colors(red, orange, yellow, purple, and white)with or without theapplication of honey water on flower models. We found that the relative strength of foragingresponse to visual versus olfactory cues differed among butterfly species. All the testedbutterflies except Danaus genutia showed a tendency to color. Cethosia cyane and Danauschrysippus mainly depended on vision to locate and search the food; Cethosia biblis, Idealeuconoe, and Tirumala limniace gave priority on olfaction to vision; Catopsilia pomona gaveequally important to olfaction and vision. We firstly found that Danaus genutia which is aspecialist to flower, mainly depended on olfaction in foraging, and was not sensitive to thetested colors. We suggest that the seven butterfly species tested exhibit four different patternsin how visual and olfactory cues were combined as determinants of foraging importance:1)Vision was given priority over olfaction;2)Olfaction was given priority over vision;3)Olfaction and vision were equally important;4)Only olfaction was used.
In butterfly foraging, olfactory signal may be the key factor, which played attractive roles;vision signal may be function as important factor in finding food sources, and then mainly relyon olfactory signal to stimulate feeding.
(2)The feeding responses of nectar-feeding butterflies to different sized flowers
All the tested butterflies favored large-sized flowers, except T. limniace. One reasonmaybe that large-sized flowers are more easily be found and produce more strongly volatiles.The other reason maybe relate to the butterflies body size. These seven butterfly speciesbelonged to middle or large sized butterflies, which need more energy to maintain their dailylife. So the larger flowers can satisfy their energy demands. In conclusion, the foragingstrategies of butterflies are not only mainly determined by the weights given to visual andolfactory cues, but also affected by body size.
(3)The visual and olfactory responses during fruit-feeding butterflies foraging
The foraging responses of foraging adults of Kallima inachus to red, yellow, purple, orwhite and to six different fermented fruits(pear, apple, banana, watermelon, orange, andpersimmon)juices were observed to evaluate which cues can be used by foraging adults. Theresults showed adults did not show foraging responses to either red, yellow, purple, or whiteartificial flowers without food odors, while flowers with the fermented pear juice stronglyattracted them, and the feeding responses to fermented juices of the six fruits show nostatistically significant different.
(4)The field behavioral and electroantennogram responses of butterflies to the foodproduced volatiles
1)The field behavioral responses of T. limniace to the volatiles of nectar-flowers
The nectar-flowers Lantana camara and Asclepias curassavica volatiles: α-pinene andmethyl salicylate, and flower-part volatiles analogues: n-capryl aldehyde, eucalyptol, andterpineol, and the common volatiles of flower: benaldehyde were used to observe the foragingresponses of T. limniace. The results showed the foraging preferences of T. limniace todifferent single compound, to mixed compounds and to different dose of α-pinene and methyl salicylate were not significantly higher than that of deionized water. Moreover, there were alsono significant differences between different compounds.
2)The field behavioral and electroantennogram responses of Kallima inachus to thefermented fruit volatiles
The foraging responses to the fermented fruits volatiles:3-methyl-1-butanol, ethyl acetate,3-methyl-1-butanoacetate, α-pinene, the2-pentanone homologue butanone, and the majorvolatiles of fermentation: ethanol, and acetic acid were observed. And the electroantennogram(EAG)responses of naive adults to these compounds were also tested. In field behavioral tests,alcohols were the most attractive, followed by esters; while α-pinene, butanone and acetic acidwere much less attractive to the butterflies. Relative to other volatile combinations and ethanolalone, the mixture of ethyl acetate and ethanol attracted the most feeding adults. The numberof adults attracted was significantly positively correlated with the concentration of bothethanol and ethyl acetate. The EAG responses of naive adults had shown that the EAGresponses to3-methyl-1-butanol,3-methyl-1-butanoacetate, ethyl acetate, α-pinene, butanoneand acetic acid were all higher than those to ethanol(100%)at doses of either5μl/ml or50μl/ml. Sexual differences only existed in3-methyl-1-butanol and acetic acid at particularconcentrations. Sexual differences in response to chemical mixtures were not significant at50μl/ml. In addition, the EAG responses in the within-sex trials were not correlated to the dosage(0.01,0.1,1,5,10, and100μl/ml)of either ethanol or ethyl acetate. The results show thatolfactory cues play a crucial role in the foraging of adult K. inachus and the foraging adultscan use a variety of chemical signals derived from food, but the feeding preference to volatileswas not necessary correlated with the EAG responses.
(5)The butterflies antennal sensilla and olfactory behavioral responses analysis
The external morphology of the antennal sensillae of four flower-visiting butterflies(D.genutia, D. chrysippus, I. leuconoe and C. pomona)and fruit-feeding butterfly(K. inachus)were examined using scanning electron microscopy. The morphology and type of antennalsensillae of males and females were the same in all species. There was no obvious sexual dimorphism within species. Five morphological sensilla types were recorded, including B hmbristles, sensilla squamiformia, two types of sensilla trichodea(types I and II), sensillacoeloconica, and sensilla chaetica. The distribution and number of sensillae were alsodescribed. The total number of sensilla was greater in males than in females of the samespecies, which may be related to courtship, in which males need more sensillae to track anddiscriminate the sex pheromones released by females. Our study suggested that sensillatrichodea II and sensilla coeloconica may play a role in detecting food odors, whereas sensillasquamiformium may serve as auxiliary sensors in food location, according to the olfactionresponses during foraging. In general, the total numbers of sensillae, or of sensilla that arerelated to food odor detection, can reflect the signal utilizing rule of foraging butterflies.
(6)The butterflies compound eyes and the preliminary analysis of foraging behavior
The microstructure and ultrastructure of the compound eyes of four flower-visitingbutterflies(D. genutia, D. chrysippus, I. leuconoe and C. pomona)and fruit-feeding butterfly(K. inachus)were examined using optical microscopy and transmission electron microscopy.There were a large number of ommatidia in five species and the total number of ommatidiawas greater in males than females of the same species. Meanwhile, there was significantdifference in the number of ommatidia in C. pomana. On the structure of compound eyes, noobvious sexual dimorphism within species was found. In general, the total number ofommatidia can basically reflect the signal utilizing rule of foraging butterflies.
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