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大多数访花昆虫对某些特定颜色、气味会表现出偏好,在寻找蜜源时,常以此作为识别、选择信号。昆虫气味物质大致包括2大类:昆虫自身产生的气味物质和植物产生的气味物质。通常,植物产生的气味物质多作为昆虫觅食、产卵的信号物质,而昆虫产生的气味物质则多被用于种群内部交流[1]。
蝶类是重要的访花传粉昆虫,在访花觅食过程中,远距离定位蜜源植物主要依靠花朵颜色提供的各种视觉信号,而嗅觉、触觉信号被认为在蜜源植物的近距离识别中发挥作用[2-4]。尤其是花色对蝴蝶搜寻蜜源相当重要,然而不同蝶种对颜色和气味的趋性有较大的区别[5-6]。目前关于蝴蝶成虫复眼的结构和功能方面已经进行过广泛的研究[7-11]。白天活动的蝴蝶光感受器多样化,不同蝶种光谱感受器种类不同[12-13]。白带锯蛱蝶(Cethosia cyane)与金斑蝶(Danaus chrysippus)在访花过程中偏爱红色、橙色和黄色[6]。而达摩凤蝶(Papilio demoleus)偏爱紫色和蓝色[14]。暗脉粉蝶(Pieris napi)在觅食时主要利用视觉[15]。除了视觉,蝴蝶还具有发达的嗅觉系统[16-17],花的气味则作为嗅觉信号影响蝴蝶的访花行为[18-19],如荨麻蛱蝶(Aglais urticae)、黄蛱蝶(Polygonia calbum)、枯叶蛱蝶(Kallima inachus)和红锯蛱蝶(Cethosia bibis)主要利用嗅觉信号进行觅食及搜寻寄主植物[6, 20-21]。
不仅如此,蝶类在求偶过程中,也需通过视觉、嗅觉和触觉等信号来进行配偶的识别[22]。其中有些蝶类以视觉为主[23-24],有些以嗅觉[25-26]或触觉为主[27-28],有的蝶类综合利用视觉和嗅觉信号[17]进行配偶识别。目前已有研究表明,暴露在蝴蝶翅膀腹面的色彩信号有助于躲避捕食者[29-30],物种识别[31],而隐藏在翅膀背面的色彩信号,包括紫外线信号,则在寻找配偶中起作用[32-33],如菜粉蝶(Pierisrapae)雌蝶对背部翅膀颜色较深的雄性表现出明显的偏好,这些雄蝶的翅膀背面颜色在长波较亮,紫外波长较暗,并且含有大量的蝶呤色素[34]。当把蝶呤从雄性翅膀上去除后,雄蝶的求偶成功率降低[35]。而在近距离求偶时,蝴蝶利用虫体挥发物提供的嗅觉信号对配偶进行精准识别,这种嗅觉信号,被认为是对蝶类视觉信号的补充[36]。成虫虫体挥发物可以确保雌雄蝶交流具有高度的私密性[20]。如雄性信息素MSP在偏瞳蔽眼蝶(Bicyclus anynana)的配偶选择中起关键作用[37-38]。
鹤顶粉蝶(Hebomoia glaucippe)又名赤顶粉蝶、红襟粉蝶,隶属于鳞翅目(Lepidoptera)粉蝶科(Pieridae)襟粉蝶族(Anthocharidini)鹤顶粉蝶属(Hebomoia)[39]。国内分布于福建、广东、海南、四川、云南、香港等地[40]。不同蝶种觅食求偶过程中的视觉嗅觉信号识别机制不同,有关鹤顶粉蝶觅食求偶过程中视觉嗅觉利用系统研究尚未见报道。本研究通过对鹤顶粉蝶觅食求偶过程中识别行为的观察,探索鹤顶粉蝶在觅食和求偶过程中的视觉和嗅觉选择,以期了解鹤顶粉蝶觅食求偶过程中对视觉嗅觉信息利用机制,为开展鹤顶粉蝶的规模化人工繁育提供理论和实用依据,为其它蝴蝶的研究提供参考。本研究亦为进一步探索蝴蝶进化过程中视觉嗅觉利用提供基础资料。
鹤顶粉蝶觅食求偶过程中的视觉嗅觉信号识别
Recognition of Visual and Olfactory Signal Functions in Hebomoia glaucippe Adults During Their Foraging and Courtship
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摘要:
目的 研究鹤顶粉蝶(Hebomoia glaucippe)成虫觅食、求偶时的视觉、嗅觉信号利用机制。 方法 通过行为观察分别测定不同花色和蜂蜜水及塑封模型对鹤顶粉蝶的引诱效应。 结果 在无气味的7种不同颜色假花诱集试验中,雌雄蝶对长波长红色(620~750 nm)表现出明显的颜色倾向,雌雄蝶访花总次数为259次,雌雄蝶分别为151次和108次。而在假花上喷洒蜂蜜水后,雌雄蝶访花总次数为583次,雌雄蝶分别为305次和278次。单一蜂蜜水诱集时,雌雄蝶访花总次数仅10次,雌雄蝶访花次数分别为8次和2次。颜色-气味组与气味组和颜色组相比,访花次数分别提高58.30倍和2.25倍。在鹤顶粉蝶求偶过程中,雌雄蝶对模型的访问和自然种群的追逐规律不一致,鹤顶粉蝶对缺乏气味的标本模型的访问次数显著低于自然种群(P<0.05)。雌雄蝶翅膀的色彩与图案存在差别。 结论 鹤顶粉蝶在觅食过程中,以视觉为主,嗅觉为辅,具有明显的色觉偏爱;在求偶过程中,视觉和嗅觉均参与求偶识别,虽然鹤顶粉蝶可以通过视觉信号进行最初的配偶识别,但仍需要通过嗅觉信号精确识别配偶及同性。 Abstract:Objective To study the mechanism of visual and olfactory signal utilization in the foraging and courtship of Hebomoia glaucippe. Method The attractive effects of different colors of flowers and artificial flowers sprayed with honey water and artificial models on H. glaucippe were verified by behavioral studies. Result In the odorless attractive experiment of 7 different colors of artificial flowers, the male and female butterflies showed an obvious color tendency towards long-wavelength red (620-750 nm). The total number of visits to flowers by the male and female butterflies was 259, 151 and 108 respectively. After spraying honey water on the artificial flowers, the male and female butterflies visited the flowers for 583 times, 305 times and 278 times respectively. In the case of using honey water only, the total number of female and male butterfly visits was only 10, in which the numbers of female and male butterfly visit were 8 and 2 respectively. Compared to the odor group and color groups, the butterfly visiting times of color + odor group increased by 58.3 and 2.25 times, respectively. During the courtship, the chasing of the model by the H. glaucippe was inconsistent with the chasing law of the natural population. The visit to the scentless specimen model was significantly lower than that of the natural population (P<0.05). There were differences in the colors and patterns of the wings of female and male butterflies. Conclusion H. glaucippe butterfly is mainly visual and supplemented by olfaction when foraging, and it has obvious preference for color vision. Both vision and olfaction participate in courtship recognition during courtship, using visual signals for initial spouse recognition, and then precise recognition through olfactory signals. -
Key words:
- Hebomoia glaucippe
- / foraging behavior
- / courtship behavior
- / visual response
- / olfactory response
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