白蚁信息素研究进展

白蚁是最古老的社会性昆虫, 其社会性的维持需要信息素的相互作用。本文回顾了近年来国内外白蚁信息素研究的最新进展, 内容涉及白蚁踪迹信息素、 性信息素、 告警信息素和促食信息素的功能、 化学成分及产生信息素的外分泌腺。白蚁分泌信息素的腺体主要有背板腺、 腹板腺、 后腹板腺、 额腺和唾腺。绝大多数白蚁信息素是挥发性物质。白蚁在化学通讯上存在节俭策略, 即同一种化合物由不同的白蚁种类的不同外分泌腺分泌, 可具有不同的功能。总结了各类信息素在白蚁物种间、 同一物种的品级间和性别间的异同和作用方式, 强调了白蚁信息素的反应阈值、 最佳浓度、 有效期和物种特异性对其功能的影响。目前对白蚁信息素的研究尚处于起步阶段, 其研究成果对等翅目系统发育研究和白蚁防治具有重要的意义。文章最后展望了白蚁信息素在白蚁防治上的应用前景。     

土垅大白蚁Macrotermes annandalei(Silvestri)采食品级的踪迹信息素

【目的】解析土垅大白蚁Macrotermes annandalei(Silvestri)采食行为中的踪迹信息交流机制。【方法】通过对其踪迹信息素进行化学和行为活性分析。【结果】土垅大白蚁兵蚁腹板腺源踪迹信息素的主要成分为顺-3-十二碳单烯醇-1((3Z)-dodec-3-en-l-ol,DOE),与工蚁的踪迹信息素化学结构完全相同。大兵蚁腺体当中DOE的含量最高,分别是大、小工蚁品级和小兵蚁品级DOE含量的3.88倍和3.78倍,与其在采食行为中的功能相适应。尽管小兵蚁的位置处于采食队伍的先锋,但小兵蚁的踪迹信息素含量与工蚁相当,不能起到召募采食的作用。行为观察和跟踪信息素行为活性表明大兵蚁具有通过分泌大量踪迹信息素召募工蚁品级的能力。【结论】土垅大白蚁依靠大兵蚁分泌大量踪迹信息素单一组分及其在采食队伍中的位置的变化来实现采食行为中的方向信息的快速传递交流。     

合成黑翅土白蚁踪迹信息素类似物的生物活性

合成了黑翅土白蚁踪迹信息素类似物(Z,Z)-3,6-十二碳二烯醇-1（DDE-OH）,该类似物对黑翅土白蚁工蚁具有和信息素提取物类似的行为反应。活性反应阈值为10^-3～10 ng/cm,大于10 ng/cm时产生较强的驱避作用。最佳活性浓度DDE-OH与信息素提取物的活性反应之间没有显著差别。     

白蚁采食行为中的信息交流

白蚁作为社会性昆虫, 其采食个体间依赖各种信息交流机制进行联系与协作, 其中包括踪迹、 警戒、 食物信息、 同伴识别和助食等。通常的联系信号为挥发性或半挥发性化学物质或者一定频谱和功率的机械振动波。其中踪迹信息素、 蚁源烃类、 警戒信息素、 助食素和机械振动信号等都在白蚁采食过程中起着重要作用。白蚁采食过程中食物定向和食物品质信息的传达主要依靠腹板腺分泌的踪迹信息素, 已发现有十二碳烯醇类、 降碳倍半萜类和大环二萜类。蚁源的机械振动也在食物品质表达方面起到一定作用, 但食物品质的表达机制还不明确。白蚁采食协作的基础是同伴识别, 蚁源烃类（C21~C35）是同伴识别的主要信息物质, 采食个体分泌的助食素则可促成共同取食。警戒信息素传递白蚁采食的安全信息, 通常为一些兵蚁源的萜类物质, 但得到功能鉴定的结构还不多。近来研究还发现特殊的蚁源机械振动也可起到示警作用。已初步证实各种信息交流机制间存在交互作用, 但交互作用的机理有待进一步解析。生物物理因素在白蚁采食行为中的作用值得更多重视。本文以白蚁的采食行为为线索, 评述白蚁采食个体间信息交流机制的研究进展及利用问题, 并对今后的研究提出展望。     

不同世代棉红铃虫性信息素的释放速率及两种组分的比例

本文采用单个活雌蛾腺体的性信息素气体收集技术,对不同世代及实验室人工饲养的CZ品系棉红铃虫Pectinophora gossypiella(Saunders)性信息素释放速率作定量分析.结果表明,各类虫源的性信息素释放速率具有明显差异;而棉红铃虫性信息素中两种组分顺,顺-7,11-十六碳二烯醇醋酸酯和顺,反-7,11-十六碳二烯醇醋酸酯的比例保持在54.8±1.2:45.2±1.2的范围;取食相同的同一代棉红铃虫的雌蛹重与其成虫的性信息素释放速率具有正相关性.     

白蚁跟踪信息素及其类似物的活性比较试验初报

自从1967年Smythe等发现某些真菌寄生过的木材中含有一种活性物质,与白蚁腹腺分泌的跟踪信息素有相似的效用以后,引起了人们探讨白蚁踪迹物质的浓厚兴趣。白蚁是一种社会性昆虫。群体间的觅食、防卫、筑巢、交哺等行为都是通过信息物质的调节来进行的。所以开展白蚁跟踪信息素及其类似物的研究,有可能为我们找出一条新的防治途径。     

松毛虫性信息素的固相微萃取及质谱和触角电位分析

松毛虫性信息素成分在腺体中含量很低。本研究利用固相微萃取（SPME）技术近距离长时间采集单头云南松毛虫Dendrolimus houi处女雌蛾释放的性信息素成分, 并进一步利用气相色谱-质谱（GC-MS）和气相色谱-触角电位联用（GC-EAD）技术分析SPME采集的性信息素成分。 结果显示: 在云南松毛虫处女雌蛾求偶召唤期间SPME聚乙二醇/二乙烯基苯（CAR/DVB）萃取头吸附了大量的反5, 顺7-十二碳二烯醇（E5, Z7-12:OH）和反5, 顺7-十二碳二烯乙酸酯（E5, Z7-12:OAc）成分及微量的反5, 顺7-十二碳二烯醛（E5, Z7-12:Ald）成分, 这3种腺体成分均能激起云南松毛虫雄虫触角电位反应。与溶剂浸提昆虫性信息素成分的方法比较, SPME活体采样技术和GC-MS及GC-EAD分析方法联用研究昆虫的信息素成分具有采集的样品代表性强、样品量能满足色谱分析需要和无溶剂干扰等方面的优点。最后还着重讨论了SPME技术的应用价值、效果及实际应用中应该注意的一些问题。      

马尾松毛虫性信息素在不同类型诱芯中的稳定性

顺5,反7-十二碳二烯醇、顺5,反7-十二碳二烯乙酸酯和顺5,反7-十二碳二烯丙酸酯是马尾松毛虫Dendrolimus punctatus性信息素的主要成分。为研究如何减少上述共轭二烯信息素成分中的几何构型异构化和氧化的发生,作者制备了几种不同剂型的诱芯并进行了田间诱蛾试验。与常用的天然橡胶诱芯相比,诱芯中加入抗氧化剂（topanol CA）、提高性信息素的纯度、每5天重新更换诱芯及将载体换为硅橡胶等处理均不能明显提高诱芯的诱蛾效果,然而载体为复合橡胶（氯化丁基橡胶和天然橡胶的混合体）的诱芯诱蛾效率比天然橡胶诱芯提高了1倍以上。田间试验后,用毛细柱气相色谱对不同类型诱芯中残留的信息素及其异构体的分析结果显示：复合橡胶诱芯中只有12%～16%的不同信息素成分发生了异构化,而在天然橡胶诱芯中此值高达69%～87%。而且前者中信息素及其异构体的剩余量是后者的4倍。这些结果表明,复合橡胶诱芯之所以具有较高的诱效,主要由于在这种诱芯中,共轭二烯信息素更稳定,释放速率较为缓慢且均匀。     

黑翅土白蚁踪迹信息素组分数量的柱层析分离和活性测定分析

本文分析了在活性限浓度进行生物活性测定时,黑翅土白蚁Odontotermes formosanu Shiraki对踪迹信息素的行为反应特点以及使用不同踪迹线载体对踪迹信息素活性的影响。并设计标准生物活性测试方法,测定了采用柱层析方法分离的黑翅土白蚁踪迹信息素各分离馏分的活性,发现黑翅土白蚁踪迹信息素用22%乙酸乙酯/正己烷洗脱液等度洗脱分离后的各馏分中包含两个强活性区段,推测黑翅土白蚁踪迹信息素至少包含2个有轻微极性差异的组分。     

黑翅土白蚁的跟踪信息素

黑翅土白蚁Odontotermes formosanus(Shiraki)严重危害堤坝,有明显跟踪行为,其分泌跟踪物质的腹板腺位于腹部第5节腹板前沿中部,分泌细胞呈长形,与腹板呈垂直方向排列。用滤纸双圈法测定黑翅土白蚁跟踪行为。跟踪物质的抽提以正己烷较好,丙酮和二氯甲烷次之。从抽提液柱层析、薄层层析和微化学反应结果推测,其跟踪物质可能是含有双键的醇类化合物。密褐褶孔菌及其感染的腐木块抽提液及白蚁跟踪信息化合物——顺,顺-3,6-十二碳双烯醇都对黑翅土白蚁有较强的跟踪活性。肖若散白蚁和家白蚁抽提液均对黑翅土白蚁有跟踪活性,但肖若散白蚁和家白蚁对黑翅土白蚁抽提液及其柱层析和薄层层析的活性馏份无活性反应。     

Nonadecadienone, a new termite trail-following pheromone identified in Glossotermes oculatus (Serritermitidae)

Within the multitude of chemical signals used by termites, the trail marking by means of pheromones is ubiquitous. Chemistry and biology of the trail-following communication have been described in more than 60 species from all families except for the Neotropical Serritermitidae. The chemical ecology of Serritermitidae is of special interest not only as a missing piece of knowledge on the diversity and evolution of isopteran pheromones but also because it may contribute to the debate on the phylogenetic position of this family, which is still unresolved. Therefore, we aimed in this study to identify the trail-following pheromone of the serritermitid Glossotermes oculatus. Based on a combined approach of analytical chemistry, electrophysiology, and behavioral bioassays, we propose (10Z,13Z)-nonadeca-10,13-dien-2-one to be the trail-following pheromone of G. oculatus, secreted by the sternal gland of pseudergates. Thus, we report on a new termite trail-following pheromone of an unexpected chemical structure, a ketone with 19 carbons, contrasting with unsaturated alcohols containing 12 carbons as trail-following pheromones in other advanced termite families. In addition to this unique trail-following pheromone, we also describe the sternal gland in pseudergates as an organ of unusual shape, size, and structure when compared with other isopteran species. These results underline the peculiarity of the family Serritermitidae and prompt our interest in the chemistry of pheromones in the other genus of the family, Serritermes.     

白蚁诱食信息素研究进展

白蚁为社会性昆虫,其组织结构和联系方式主要由白蚁外分泌腺产生的起诱导和调节白蚁行为反应作用的信息素来获得。由白蚁下唇腺产生的诱食信息素,能够在白蚁巢体的群体性食物采集中诱使取食白蚁形成聚集,并且取食食物的相同位置,从而提高白蚁巢体采集食物的效率。目前,已确定对苯二酚为白蚁诱食信息素,且认为整个等翅目昆虫都产生和使用对苯二酚作为诱食信息素,与分类地位和生物学特性无关。该文概述了白蚁诱食信息素的分泌器官、种特异性、生物学意义、生物合成途径及其在白蚁防治中的应用等方面的研究进展。     

Identification of multi‐component trail pheromones in the most evolutionarily derived termites,the Nasutitermitinae (Termitidae)

In the present study, trail pheromone blends are identified for the first time in termites. In the phylogenetically complex Nasutitermitinae, trail‐following pheromones are composed of dodecatrienol and neocembrene, the proportions of which vary according to species, although neocembrene is always more abundant than dodecatrienol (by 25–250‐fold). Depending on species, termites were more sensitive to dodecatrienol or to neocembrene but the association of both components always elicited significantly higher trail following, with a clear synergistic effect in most of the studied species. A third component, trinervitatriene, was identified in the sternal gland secretion of several species, but its function remains unknown. The secretion of trail pheromone blends appears to be an important step in the evolution of chemical communication in termites. The pheromone optimizes foraging, and promotes their ecological success. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 20–27.     

Interaction of pheromones during food exploitation by the termite Schedorhinotermes lamanianus

Abstract. Chemical signals from secretions of different exocrine glands modulate a variety of behavioural patterns in termite societies. These signals have multiple functions and may be interactive. During food exploitation workers of the African termite Schedorhinotermes lamanianus (Isoptera: Rhinotermitidae) employ, on foraging trails, the secretion from the sternal gland both for orientation and recruitment to a food source. The secretion from the labial gland, released onto the food by gnawing termites, stimulates additional workers to gnaw at the same site, thereby forming aggregations of gnawing termites. An interaction between these two pheromones during food exploitation is demonstrated for the first time. The volatile signal from the sternal gland inhibits in a dose-dependent manner the non-volatile, highly persistent, signal from the labial gland. The development of gnawing aggregations is inhibited and established ones are dissolved. Behavioural evidence for the perception of both the volatile signal from the sternal gland by olfactory neurones and of the non-volatile signal from the labial gland by gustatory neurones on the antennae is given. The interaction of the two pheromones as a basis for the development of distinct commuting and gnawing zones on the food source, and as a means for a dynamic regulation of food exploitation, is discussed.     

Species-specific sex pheromones secreted from new sexual glands in two sympatric fungus-growing termites from northern Vietnam, <Emphasis Type="Italic">Macrotermes annandalei</Emphasis> and <Emphasis Type="Italic">M. barneyi</Emphasis>

Summary Reproductive isolation in termites is not well known. Our study carried out on two sympatric species from northern Vietnam, Macrotermes annandalei and M. barneyi, showed that dispersal flights and sex pheromones were two important factors in their reproductive isolation. These fungus-growing termites were isolated, partially due to the timing of their respective dispersal flights. M. annandalei flew the first day after rain, while the flights of M. barneyi occurred the second day after rain. However, the flights can also be simultaneous in the two species. Sex pheromones of M. annandalei and M. barneyi were shown to be species-specific. In both species, they were secreted by females from two glandular sources, from tergal glands located on tergite 6 to 10 in M. annandalei and tergite 5 to 10 in M. barneyi, and from posterior sternal glands located on sternite 6 and 7 in both species. These posterior sternal glands, found for the first time in the Termitidae, were sex-specific glands. Although not fully identified, sex pheromones of M. annandalei and M. barneyi were clearly different from the trail-following pheromone secreted by the sternal gland stricto sensu located on the sternite 5. These results show that in termites, the sexual behaviour, the glandular origin of sex pheromones and their role in reproductive isolation greatly vary depending on the species and deserve to be more extensively studied.Received 8 April 2003; revised 1 September 2003; accepted 10 September 2003.     

Trail Communication During Foraging and Recruitment in the Subterranean Termite Reticulitermes santonensis De Feytaud (Isoptera, Rhinotermitidae)

The search for food in the French subterranean termite Reticulitermes santonensis De Feytaud is organized in part by chemical trails laid with the secretion of their abdominal sternal gland. Trail-laying and -following behavior of R. santonensis was investigated in bioassays. During foraging for food termites walk slowly (on average, 2.3 mm/s) and lay a dotted trail by dabbing the abdomen at intervals on the ground. When food is discovered they return at a quick pace (on average, 8.9 mm/s) to the nest, laying a trail for recruiting nestmates to the food source. While laying this recruitment trail the workers drag the abdomen continuously on the ground. The recruitment trail is highly attractive: it is followed within a few seconds, by more nestmates, and at a quicker pace (on average, 6.4 mm/s) than foraging trails (on average, 2.9 mm/s). The difference between foraging and recruitment trails in R. santonensis could be attributed to different quantities of trail pheromone. A caste-specific difference in trail pheromone thresholds, with workers of R. santonensis being more sensitive to trails than soldiers, was also documented: soldiers respond only to trails with a high concentration of trail pheromone.     

The role of pheromones in the initiation of foraging, recruitment and defence by the soldiers of a tropical termite, Nasutitermes corniger (Motschulsky)

The polyethic behaviour of soldiers and workers of Nasutitermescorniger during foraging and in defence is described. It hasbeen shown that the cephalic gland defensive secretion of thenasute soldiers produces short term recruitment of only soldiersto loci where it is present. Without further reinforcement ofthe stimulus, the number of soldiers wanes. Workers react tothe cephalic gland secretion by remaining in or retreating tothe nest. The sternal gland secretion of workers and soldiersacts in long term recruitment of soldiers when presented experimentallyas a point source, as a trail or as a trail with a food sourceat its end. This secretion causes more exiting soldiers thanthe cephalic gland secretion, and is more effective in thisregard as a trail than as a point source. The greater numbersexiting in the trail situation appears to be related to thegreater amount of pheromone area present, and the halting ofsoldiers at the pheromone odour boundary. The greatest numberof soldiers exit when the trail is coupled with a food sourceat its end, and in this situation workers exit for the firsttime in significant numbers. Additional information about thefood source is postulated to be communicated by tactile means(jittering) and it is suggested that this lowers the thresholdfor the trail following responses in the termites such thatworker exiting is initiated. Other factors affecting the numbersforaging are noted. The phenomenon of soldier initiation offoraging as scouts, and as a scouting column, is commented onwith regard to the polyethic responses to the pheromones, andto its adaptive significance (sociobiological import) in thelight of the soldiers' rôle in colony defence.     

Fire ant trail pheromones: Analysis of species specificity after gas chromatographic fractionation

The specificity of the trail pheromones of four Solenopsis species was determined using natural trails. Dufour's gland extracts, and purified fractions from Dufour's gland extracts collected after gas-chromatographic separation. S. richteri and S. invicta possess species-specific major trail pheromones, while S. geminata and S. xyloni appear to have a common trail pheromone. Preliminary chemical characterization of the main trail pheromone of S. richteri indicates a M.W. of 218 and empirical formula of C₁₆H₂₆. The trail pheromone system of S. richteri consists of a blend of compounds and this phenomenon may also occur in the other species. The lowest concentration of their trail pheromone that workers of S. richteri could detect was about 10 fg per cm. The significance of blends of pheromones being utilized to generate chemical trails is discussed.