摘要
在漫长的进化历程中,植物花的结构不断变得复杂,颜色不断变得鲜艳,花中具有花蜜和花粉,大多数花具有香味,这些特征使植物向着更有利于招引昆虫的方向发展,而昆虫的形态结构和生活习性也相应地发生变化,向着便于采蜜传粉的方向发展。从而,在植物和昆虫之间产生了精巧而微妙的相互适应和协同进化关系。本论文以长白山地区为研究范围,对主要蜜源植物与四类重要访花昆虫类群之间的相互适应关系进行了研究,包括:主要蜜源植物花部综合特征(花的结构、颜色、花粉形态及花的气味)与重要访花昆虫类群的适应关系;重要访花昆虫类群外部形态(口器、携粉足等)、内部系统(消化道的形态结构)、微观结构(触角感受器的类型、数量、分布规律)与昆虫访花行为的适应关系。本研究可为开发新型昆虫引诱剂及授粉昆虫在设施农业领域中的应用提供了重要的理论参考。研究所采用的主要方法及结果如下:
(1)对长白山地区12个市、县、镇及长白山自然保护区共计22个样地内的访花昆虫进行了野外采集、标本鉴定和文献检索,共获得访花昆虫452种,隶属于10目、74科、283属。其中,膜翅目蜜蜂科(蜜蜂属和熊蜂属)和隧蜂科昆虫、双翅目食蚜蝇科昆虫、鞘翅目天牛科(花天牛亚科)昆虫和鳞翅目蝶类昆虫为长白山地区重要访花昆虫类群。
(2)对长白山地区主要蜜源植物的访花昆虫类群调查显示,菊科植物的主要访花昆虫类群为膜翅目、双翅目和鳞翅目蝶类昆虫;蔷薇科植物的主要访花昆虫类群为膜翅目熊蜂属、鞘翅目花天牛亚科和鳞翅目蝶类昆虫;毛茛科、唇形科、豆科植物的主要访花昆虫类群为膜翅目昆虫;百合科、石竹科植物的访花昆虫类群为鳞翅目蝶类昆虫;伞形科植物的主要访花昆虫类群为膜翅目熊蜂属和鞘翅目花天牛亚科昆虫;败酱科植物的主要访花昆虫类群为双翅目昆虫;蓼科、罂粟科和牻牛儿苗科植物的主要访花昆虫类群为膜翅目和双翅目昆虫;柳叶菜科植物的主要访花昆虫类群为膜翅目和鳞翅目蝶类昆虫。
(3)对四类重要访花昆虫偏爱的蜜源植物的花朵形态及颜色分析显示,熊蜂偏爱访问具有圆锥花序、总状花序和聚伞花序的花;花冠类型以辐射对称、两侧对称为主;花色艳丽,常为黄色或蓝紫色。天牛访问的花多为圆锥、伞形、聚伞和伞房花序;花冠类型以辐射对称为主,多为旋转状或坛状,而非管状或唇形花;具有大量的雄蕊和花粉,通常缺少花蜜;由于天牛通常缺乏对颜色的洞察力,主要访问白色和黄色等淡色花,很少访问红色或紫色花。食蚜蝇偏爱访问具有头状花序和伞形花序的花朵;由于其喙管较长,可以访问筒形花,吸食花基部的蜜汁;对花色的选择能力较强,但多访问白色和黄色的花朵。蝶类偏爱访问具有头状花序和总状花序,色泽鲜艳的花朵。
(4)利用扫描电子显微镜观察了长白山地区主要蜜源植物的花粉形态,并分析了花粉形态与四类重要访花昆虫类群的关系。结果表明,熊蜂属昆虫和花天牛亚科昆虫偏爱访问的蜜源植物均具有长球形花粉,花粉粒较大,P/E值大于1.14,具三孔沟,表面具有网状纹饰。食蚜蝇科和蝶类昆虫对蜜源植物的花粉要求不高。
(5)利用气相色谱-质谱联用仪分析了长白山地区11种主要蜜源植物的乙醇提取物成分,并对其进行了比较。结果表明,蜜源植物花乙醇提取物成分在菊科、蔷薇科、伞形科、柳叶菜科和豆科之间未表现出显著的特异性,主要物质为酸类化合物,其次是酮类化合物和酯类化合物。11种花乙醇提取物成分中均含有1–羟基–2–丙酮、乙酸、甲酸、顺式–9,12–十八碳二烯酸和苯酚,且这些成分的含量分别在提取物总含量中具有相似的比例关系。可以推测这些成分对访花昆虫具有引诱作用。但对昆虫产生引诱性的是这些混合物中各成分的综合效果,还是其中的某一种物质起着主要的引诱作用,还有待于进一步的研究。
(6)通过对访花昆虫类群外部形态特征与访花的适应研究显示:①熊蜂较为粗壮;全身密被长而整齐的毛,能粘有大量花粉;嚼吸式口器,中唇舌较长,能从深花冠花的正中吸食花蜜;具有花粉筐,后足基胫节内表面具毛刷,能够携带大量花粉。②复眼通常较大并有很多小眼,使其能顺利地搜索到所需的寄主植物;翅发达且常有色,触角相对较长,通常丝状或锯齿状,多数位于头部顶端而非侧面,为花天牛携带花粉进行大范围的转移提供了可能。③食蚜蝇类可通过体毛粘附花粉;舐吸式口器能分泌涎液稀释浆状的花蜜以便自身的吸收;跗节具有化学感受器,对含糖液汁敏感。④蝶类为虹吸式口器,善于吸食花蜜;具有较发达的足,访花时常抓附于花上,粘有大量的花粉;视觉发达,倾向于访问色泽鲜艳的花朵。对于访花昆虫适于访花特征的研究,将有利于挖掘新的授粉昆虫。
(7)熊蜂属昆虫、花天牛亚科昆虫、食蚜蝇科昆虫和蝶类昆虫的消化道均由前肠、中肠和后肠组成。熊蜂属昆虫的蜜囊明显,内壁形成纵横的褶皱,容积较大,能够容纳大量蜜汁。中肠较粗,内壁由多条纵嵴组成,在嵴上有许多突起,能够增加中肠的表面积,有助于中肠消化液的分泌和对营养物质的吸收。花天牛亚科昆虫取食花粉和花瓣,嗉囊和中肠的内壁形成了许多乳状的盲囊。食蚜蝇科昆虫和蝶类昆虫主要取食花蜜等液体食物,其嗉囊内壁和中肠内壁较为平滑,主要起吸收的作用。
(8)熊蜂属昆虫触角上具有毛形感受器(A、B、C/ D)、锥形感受器(A、B)、腔锥形感受器、板形感受器和Bohm’s鬃毛;花天牛亚科昆虫触角上具有毛形感受器(Ⅰ、Ⅱ、Ⅲ)、刺形感受器(Ⅰ、Ⅱ、Ⅲ、Ⅳ)、锥形感受器(Ⅰ、Ⅱ、Ⅲ)和Bohm’s鬃毛;食蚜蝇科昆虫触角上具有毛形感受器(Ⅰ、Ⅱ)和大量微毛;蝶类昆虫触角上具有鳞形感受器、星形感受器、毛形感受器(A、B)。毛形感受器为四类访花昆虫类群触角上共有的感受器类型,通常作为机械力感受器,因而可以推断其对于访花昆虫在花朵上的栖息起到重要的作用。此外,四类访花昆虫类群还各具有特异的感受器类型,推断其对识别特异花朵具有重要的意义。基于访花昆虫与植物的紧密联系,为了阐明不同类群嗅觉感受器的功能及作用,还需进一步对触角感受细胞的电生理学进行研究。
In the long course of evolution, flowers and insects produced a sophisticated and subtle relation of mutual adaptation. The structure of flower becomes increasingly complex, the flowers clustered, the color of flower becomes increasingly bright, has nectary, pollen and fragrance, toward the direction of more conducive to attract insects. And, accordingly, morphology and living habits of insects have also changed, towards the direction of collecting nectar and pollination. This study considered the mutual adaptation between the main nectar plants and flower-visiting insects in Changbai Mountain Region through the floral features, including flower structure, color, pollen morphology, odor, and insects’characteristics, including the external morphology of insects, such as the mouthpart, portability foot powder; the morphology and structure of alimentary canal, and the type, number and distribution of antennal sensilla. This study not only helps on the rare nectar plants and ecosystem protection, but also for the development of new insect attractant, and provides a theoretical basis for insect pollination in the field of facilities agriculture. The main methods used in this study and the findings are as follows:
(1) A total of 452 species of flower-visiting insects in Changbai Mountain Region were collected, indentified and literature searched, which belonging to 10 orders,74 families and 283 genera.
(2) The major visitors of Compositae flowers were Hymenoptera insects, Diptera insects, and Lepidoptera butterflies; the major visitors of Rosaceae flowers were Hymenoptera bumblebees, Lepidoptera butterflies and Coleoptera flower longhorn beetles; the major visitors of Ranunculaceae flowers, Legumes flowers and Labiatae flowers were Hymenoptera insects; the major flower-visiting insects of Hyacinth and Caryophyllaceae were Lepidoptera butterflies; the major visitors of Umbelliferae flowers were Hymenoptera bumblebees, Coleoptera flower longhorn beetles; the major visitors of Polygonum flowers, Poppy flowers, Geraniaceae flowers were Hymenoptera and Diptera insects; the major visitors of Valerianaceae flowers were Diptera insects; the major visitors of Onagraceae flowers were Hymenoptera and Lepidoptera butterflies.
(3) Bumblebees prefered panicle, raceme and cyme, and mainly visited flowers with corolla type of radial symmetry and bilateral symmetry. In the sight of flower color, bumblebees visited colorful flowers, often yellow or blue-purple. Lepturinae visited larger–shaped flowers, mainly panicles, umbel, cyme and corymb. Lepturinae feed on pollen and petal, so they visited flowers with a large number of stamens and pollen, and often lack nectar. Lepturinae visited flowers mainly with corolla type of radial symmetry, and the shape of flowers were rotating, altar-like, rather than tubular or lipped. In the sight of flower color, Lepturinae often visited white, pale yellow flowers, rarely with red or purple because of lacking color vision. Syrphidae like to visit Asteraceae and Umbelliferae plants. In the sight of Inflorescence types, Syrphidae prefered capitulum and umbel. Because of their long beak tube, they can visit the tube-shaped flowers, taking the honey from the base of flower. Syrphidae had stronger ability in the choice of colors, they prefered to visit white and yellow flowers. Butterflies prefered the flower with capitulum and raceme, and tend to visit the brightly-colored flowers.
(4) In this study, the pollen morphology of the main nectar plants in Changbai Mountain region were observed using scanning electron microscopy, and the relationship between pollen morphology and main flower-visiting insects were analyzed. The results showed that Bombus and Lepturinae perfered prolate, larger pollen (P/E values are greater than 1.14) with three–hole groove and reticulate surface. Syrphidae and butterflies were not strict with pollen morphology.
(5) The composition of ethanol extracts from 11 major nectar plants in Changbai Mountain area were analyzed using gas chromatography-mass spectrometer and compared in this study. The results showed that it didn’t had signifiant different among the components of ethanol extract from Asteraceae, Rosaceae, Umbelliferae, Onagraceae and legumes. The main substance were acids, followed by ketones and esters. The ethanol extracts from 11 major nectar plants were all containing 1 - hydroxy -2 - acetone, acetic acid, formic acid, cis - 9, 12 - octadecadienoic acid and phenol. These five kinds of component had a similar proportional relationship, among them, 1 - hydroxy - 2 - acetone, cis - 9, 12 - octadecadienoic acid accounted for about 20%, acetic acid accounted for about 40%, formic acid and phenol were about 10%, respectively. Presumably, these components have a lure on the attraction of flower-visiting insects. But, if it is produced by the mixture or by on of these substance is need to be further study.
(6) The adaptation on the external morphological feature of insects for visiting flower:①Bumblebees: more sturdy; with hairs, casued a large number of pollen stickied in the process of visiting; chewing-lapping mouthparts, longer tongue, making them take nectar from flowers with deep corolla; wide hind tarsus, with smooth surface and long-hairs around the end, forming pollen baskets; had wide hind coxal and tibia, with neatly arranged brushes in the inner surface, making them carry large amounts of pollen.②Lepturinae: front of the head tend to elongate, sometimes forming a beak-like protuberances, and a number of bristles and the rounded prominent part in the upper lips (suitable for lick nectary); usually have larger compound eyes and many small eyes, so that beetles can successfully search for the host plants; wings well developed and often color; antennae relatively long, usually silk or jagged, mostly located in the head rather than side, which makes beetles adapt to regular flights, and provides the possibility for beetles to carry pollen to a wide range.③Syrphidae: can adhere pollen through the body hairs; Licking mouthparts can secrete saliva diluted slurry in order to absorb the nectar; Tarsus with chemical receptors which sensitive to the sugar.④Butterflies: Siphoning mouthpa are good at feeding nectar; have more developed feet, which grasp to the flowers, and can adhere a large number of pollen; Developed vision, tend to visit brightly–colored flowers. The study on the characteristic of insects which suitable for visiting flowers will be conductive to tap the new pollinators.
(7) Alimentary canal of Bombus, Lepturinae, Syrphidae and butterflies are all composed of foregut, midgut and hindgut. For Bombus, honey sac was obvious, with vertical and horizontal folds on its wall, and larger to accommodate honey. Midgut thick, inner wall formed by a number of longitudinal ridges, in which there were many processes that can increase the intestinal surface area, and helping in the intestinal secretion of digestive juice and nutrient absorption. Lepturinae insect feed on pollen and flower petals, the inner wall of its crops and midgut’s inner formed a number of milky blind sac. Syrphidae insects and butterflies feed on nectar and other liquid food, its crop wall and intestinal wall rather than smooth, playing a major role in absorption.
(8) Sensilla trichodea (A, B, C/ D), sensilla basiconica (A, B), sensilla coeloconica, sensilla placodea and Bohm’s bristles were differentiated according to their size and shape on the antennae of Bombus; Sensilla trichodea (Ⅰ,Ⅱ,Ⅲ), sensilla chaetica (Ⅰ,Ⅱ,Ⅲ,Ⅳ), sensilla basiconica (Ⅰ,Ⅱ,Ⅲ) and Bohm’s bristles were observed on the antennae of Lepturinae; Sensilla trichodea (Ⅰ,Ⅱ) and microtrichia were on the antennae of Syrphidae; And sensilla trichodea (A, B), sensilla squamiformia and sensilla stellate were on the antennae of butterflies. There were sensilla trichodea inferred as one important receptor on all this four insect groups, which play an important role in the habitat of flower-visiting insects. Besides, each group had its own type of sensilla. Of the close relationship between flower–visiting insects and plants in view, in order to clarify the function of the olfactory receptors of different kinds of insects, the further study on the antennal cell electrophysiological are needed.
引文
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