甲虫表皮碳氢化合物研究进展

昆虫表皮碳氢化合物(cuticular hydrocarbons)是正烷烃、不饱和烃以及甲基支链烃的混合物.除了保持水分的基本功能外,还具有多种生物功能.鞘翅目昆虫(俗称甲虫)是多样性最为丰富的昆虫类群,目前对其表皮碳氢化合物研究开展颇多,主要见于化学通讯领域,内容涉及到性信息素、标记信息素、聚集信息素、化学拟态、寄主...     

Low level of cuticular hydrocarbons in a parasitoid of a solitary digger wasp and its potential for concealment

Insect cuticular hydrocarbons (CHC) play a role as semiochemicals in many host–parasite systems and chemical mimicry or camouflage is a well-known mechanism of parasites to evade detection by the host. The cuckoo wasp Hedychrum rutilans (Hymenoptera, Chrysididae) is a parasitoid of larvae of the European beewolf Philanthus triangulum (Hymenoptera, Crabronidae). Females chemically mimic the cuticular hydrocarbons of their hosts to avoid detection and countermeasures when entering the host nest for oviposition. Here we report on a possible second mechanism of the chrysidid wasp H. rutilans to evade detection: the amount of CHC/mm² of cuticle is only approximately one-fifth compared to its beewolf host. Furthermore, we show that surprisingly large amounts of CHC of beewolf females can be found on the walls of the underground nest. Potentially, these hydrocarbons might constitute a background odor against which the cuckoo wasps or their chemical traces have to be perceived by the beewolf. The reduction in the amount of CHC of the cuckoo wasps might be equivalent to a dilution of recognition cues, especially against the background odor of the nest walls, and might provide a means to escape detection within the nest due to "chemical insignificance".     

水椰八角铁甲不同日龄和性别成虫表皮碳氢化合物的种类及动态

水椰八角铁甲Octodonta nipae(Maulik)是一种严重为害棕榈科植物的外来入侵害虫。为了明确日龄和性别等个体因素对水椰八角铁甲成虫表皮碳氢化合物的影响,本研究采用化学萃取法和气相色谱-质谱联用技术,分析了不同日龄的水椰八角铁甲雌、雄成虫表皮碳氢化合物的组成和含量。结果表明,水椰八角铁甲成虫的表皮碳氢化合物由21种C_(18)-C_(30)的碳氢化合物构成,包括11种正构烷烃、2种单甲基烷烃、1种多甲基烷烃、3种正构烯烃、1种单甲基烯烃和3种环烷烃。不同日龄和性别成虫的表皮碳氢化合物的组成种类相同,但是一些组分的含量有明显变化,例如1 d、5 d、10 d和15 d雌、雄成虫之间正二十四烷、正二十六烷、2-甲基十九烷和1-二十烯的含量存在显著差异。水椰八角铁甲成虫表皮碳氢化合物具有性二型,如性成熟雄虫的表皮的正二十烷、正二十一烷、正二十四烷、正二十六烷、2-甲基十九烷、10-甲基二十烷、1-二十烯和1-二十二烯的含量显著高于性成熟雌虫。本文的研究结果可为检验检疫过程中水椰八角铁甲近缘种,以及日龄、性别和性成熟的快速鉴定提供支持。     

A chemical level in the coevolutionary arms race between an ant social parasite and its hosts

Here we investigate the coevolutionary interactions between the slavemaking ant Protomognathus americanus and its Temnothorax hosts on a chemical level. We show that, although this social parasite is principally well-adapted to its hosts' cuticular hydrocarbon profile, there are pronounced differences in the fine-tuning of this adaptation. Between populations, chemical adaptation varies with host community composition, as the parasite faces a trade-off when confronted with more than one host species. In addition to adaptation of its own chemical signature, the slavemaker causes a reciprocal adjustment in its slaves' cuticular profile, the degree of which depends on the slave species. On the host side, successful parasite defence requires efficient enemy recognition, and in behavioural aggression trials, host colonies could indeed discriminate between invading slaves, which commonly accompany slavemakers on raids, and free-living conspecifics. Furthermore, hosts shifted their acceptance threshold over the seasons, presumably to reduce the costs of defence.     

Learning and discrimination of cuticular hydrocarbons in a social insect

Social insect cuticular hydrocarbon (CHC) mixtures are among the most complex chemical cues known and are important in nest-mate, caste and species recognition. Despite our growing knowledge of the nature of these cues, we have very little insight into how social insects actually perceive and discriminate among these chemicals. In this study, we use the newly developed technique of differential olfactory conditioning to pure, custom-designed synthetic colony odours to analyse signal discrimination in Argentine ants, Linepithema humile. Our results show that tri-methyl alkanes are more easily learned than single-methyl or straight-chain alkanes. In addition, we reveal that Argentine ants can discriminate between hydrocarbons with different branching patterns and the same chain length, but not always between hydrocarbons with the same branching patterns but different chain length. Our data thus show that biochemical characteristics influence those compounds that ants can discriminate between, and which thus potentially play a role in chemical signalling and nest-mate recognition.     

Experimental and phylogenetic evidence for floral mimicry in a guild of fly-pollinated plants

Mimicry, as an adaptive explanation for the resemblance between organisms, is not always readily distinguishable from, inter alia , coincidence, shared ancestry, or convergent evolution. We tested the hypothesis that two rare South African orchid taxa Brownleea galpinii ssp. major (nectar-producing) and Disa cephalotes ssp. cephalotes (non-rewarding) are mimics of the nectar-producing flowers of a relatively common species, Scabiosa columbaria (Dipsacaceae), with which they always occur sympatrically. Flowers of the orchids were apparently unscented and had similar dimensions and almost identical spectral reflectance to the flowers of Scabiosa . The orchids were pollinated exclusively by long-proboscid flies (Tabanidae and Nemestrinidae) that feed mainly on nectar in Scabiosa flowers. Choice experiments showed that these flies did not discriminate between the orchids and Scabiosa when alighting on their flat-topped inflorescences. However, flies were not attracted to related orchids dissimilar to Scabiosa , or to inflorescences of B. galpinii that had been artificially reconstructed in the shape of a spike, rather than a flat-topped capitulum. A phylogenetic analysis showed traits that give the orchids a resemblance to Scabiosa , such as a flat-topped inflorescence and cream floral colouration with dark spots and short spurs, to be mostly apomorphic features, and therefore likely to be relatively recent adaptations for mimicry. We caution that the term mimic should not be applied to species whose resemblance to another species is due entirely to plesiomorphic traits that, in all likelihood, evolved prior to the ecological association.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 289–304.     

Sex pheromone mimicry in the early spider orchid (Ophrys sphegodes): patterns of hydrocarbons as the key mechanism for pollination by sexual deception

We investigated the female-produced sex pheromone of the solitary bee Andrena nigroaenea and compared it with floral scent of the sexually deceptive orchid Ophrys sphegodes which is pollinated by Andrena nigroaenea males. We identified physiologically and behaviorally active compounds by gas chromatography with electroantennographic detection, gas chromatography-mass spectrometry, and behavioral tests in the field. Dummies scented with cuticle extracts of virgin females or of O.sphegodes labellum extracts elicited significantly more male reactions than odorless dummies. Therefore, copulation behavior eliciting semiochemicals are located on the surface of the females' cuticle and the surface of the flowers. Within bee and orchid samples, n-alkanes and n-alkenes, aldehydes, esters, all-trans-farnesol and all-trans-farnesyl hexanoate triggered electroantennographic responses in male antennae. Most of the alkanes and alkenes occurred in similar patterns both in the bees and orchids. O. sphegodes leaf extracts contained mostly the same compounds but in different proportions. In behavioral tests with synthetic compounds, blends of alkenes triggered significantly more approaches and pounces of the males whereas alkanes were not more attractive than odorless dummies. Since alkanes and alkenes together were most attractive, we conclude they constitute the bees' sex pheromone as well as the pseudocopulation-behavior releasing orchid-odor bouquet. Accepted: 30 March 2000     

Ecology of a bolas spider,Mastophora hutchinsoni: phenology,hunting tactics,and evidence for aggressive chemical mimicry

Summary Bolas spiders are relatively rare members of the large family known as orb weavers. Instead of using a typical web to capture prey, late-stadia and adult female bolas spiders swing a droplet of adhesive on a thread at flying insects. Mastophora hutchinsoni (Araneae: Araneidae) is one of five Mastophora species known from the United States and occurs over much of eastern North America. It is univoltine in Kentucky and overwinters in the egg stage. Spiderling emerged in May, the diminutive males matured in late June and early July, and females matured in early September. Eggs were produced from late September to late October or early November. This report is the first complete documentation of the population phenology of any bolas spider. Newly-emerged M. hutchinsoni spiderlings did not use a bolas, but instead hunted by positioning themselves on the underside of leaf margins where they ambushed small arthropods that crawled along the leaf margins. Subadult and adult female M. hutchinsoni used a bolas to capture moths. Only male moths were captured, specifically three species of Noctuidae (bristly cutworm, bronzed cutworm, and smoky tetanolita) and one species of Pyralidae (bluegrass webworm). Among 492 prey captured by more than twenty spiders at two sites during 1985 and 1986, smoky tetanolita moths and bristly cutworm moths accounted for 93% of the total. The flight behavior of approaching moths, the limited taxa caught from a large available moth fauna, and the fact that only males were caught support the hypothesis that the spider attracts its prey by producing chemicals which mimic the sex pheromones of these moth species. Adult female M. hutchinsoni frequently captured more than one moth species on a given night. The two most common prey species were active at different times of night, the bristly cutworm soon after nightfall and the smoky tetanolita generally between 11:00 p.m. and dawn. This pattern suggests that mating activity of these moth species may be temporally isolated, a common phenomenon when sympatric species have similar pheromones. If so, the spider could capture both species without producing different pheromone-mimicking compounds, simply by hunting during the activity period of each species.The investigation reported in this paper (No. 87-7-76) is in connection with a project of the Kentucky Agricultural Experiment Station and is published with the approval of the Director     

Polarization sensitivity as a contrast enhancer in pelagic predators: lessons from in situ polarization imaging of transparent zooplankton

Because light in the pelagic environment is partially polarized, it has been suggested that the polarization sensitivity found in certain pelagic species may serve to enhance the contrast of their transparent zooplankton prey. We examined its potential during cruises in the Gulf of Mexico and Atlantic Ocean and at a field station on the Great Barrier Reef. First, we collected various species of transparent zooplankton and micronekton and photographed them between crossed polarizers. Many groups, particularly the cephalopods, pelagic snails, salps and ctenophores, were found to have ciliary, muscular or connective tissues with striking birefringence. In situ polarization imagery of the same species showed that, while the degree of underwater polarization was fairly high (approx. 30% in horizontal lines of sight), tissue birefringence played little to no role in increasing visibility. This is most likely due to the low radiance of the horizontal background light when compared with the downwelling irradiance. In fact, the dominant radiance and polarization contrasts are due to unpolarized downwelling light that has been scattered from the animal viewed against the darker and polarized horizontal background light. We show that relatively simple algorithms can use this negative polarization contrast to increase visibility substantially.     

Context‐dependence in an ant–aphid mutualism: direct effects of tending intensity on aphid performance

1. In ant–hemipteran mutualisms, ants receive carbohydrates in the form of honeydew, while hemipterans receive protection from natural enemies. In the absence of natural enemies, however, the direct effects of tending are generally less well known. We hypothesised that with increasing tending intensity (ant to aphid ratio), aphid performance would increase initially, then decrease at high tending levels due to the metabolic cost of producing high quality honeydew. 2. We tested our hypothesis in a greenhouse experiment by manipulating Argentine ant (Linepithema humile Mayr) colony size while holding constant the initial size of aphid (Chaitophorus populicola Thomas) aggregations. The two parameters associated with survival, aphid survivorship to maturity and longevity, declined with increasing tending intensity, whereas per capita birth rate and time to first reproduction showed no relationship to attendance. The intrinsic rate of increase declined only at relatively high tending levels, suggesting a nonlinearity in the effect of tending intensity. 3. Tending intensity measured in the experiment was similar to that observed in free‐living aggregations of C. populicola. Furthermore, the per capita recruitment rate of ants to free‐living aphid aggregations was negatively density‐dependent, indicating that small aggregations tend to experience the highest levels of tending intensity. This finding suggests that the aphid's intrinsic rate of increase may be positively density‐dependent, mediated by the aphid's mutualistic interaction with the ant. 4. In the Argentine ant–C. populicola interaction, experimental manipulation of colony size revealed a direct cost of ant attendance that was conditional upon tending intensity. Experiments that manipulate only ant presence or absence may yield an incomplete understanding of the mutualistic interaction if underlying nonlinearities exist.     

Size-mediated response to public cues of predation risk in a tropical stream fish

In order to investigate any size-dependent differences between behavioural patterns, wild-caught Hart's rivulus Rivulus hartii of varying sizes were exposed to chemical alarm cues extracted from the skin of conspecifics or heterospecific Poecilia reticulata, or a tank water control, in a series of laboratory trials. In response to conspecific alarm cues, R. hartii subjects of the range of body sizes tested exhibited consistent, size-independent antipredator behaviours that were characterized by decreased locomotory activity and foraging levels and increased refuging behaviour. Conversely, focal R. hartii demonstrated significant size-dependent trends in response to heterospecific alarm cues, with smaller individuals exhibiting antipredator responses and larger individuals shifting their behaviour to increased levels of activity consistent with a foraging, or predatory, response. These results show that the behavioural responses of individual R. hartii to publicly available chemical alarm cues from heterospecifics are mediated by the size of the receiver.     

Characterization of beta-turn and Asx-turns mimicry in a model peptide: stabilization via C--H . . . O interaction

The chemical synthesis and single-crystal X-ray diffraction analysis of a model peptide, Boc-Thr-Thr-NH2 (1) comprised of proteinogenic residues bearing an amphiphilic Cbeta -stereogenic center, has been described. Interestingly, the analysis of its molecular structure revealed the existence of a distinct conformation that mimics a typical beta-turn and Asx-turns, i.e., the two Thr residues occupy the left- and right-corner positions. The main-chain torsion angles of the N- and C-terminal residues i.e., semiextended: phi = -68.9 degrees , psi = 128.6 degrees ; semifolded: phi = -138.1 degrees , psi = 2.5 degrees conformations, respectively, in conjunction with a gauche- disposition of the obligatory C-terminus Thr CgammaH3 group, characterize the occurrence of the newly described beta-turn- and Asx-turns-like topology. The preferred molecular structure is suggested to be stabilized by an effective nonconventional main-chain to side-chain Ci=O . . . H--Cgamma(i+2)-type intraturn hydrogen bond. Noteworthy, the observed topology of the resulting 10-membered hydrogen-bonded ring is essentially similar to the one perceived for a classical beta-turn and the Asx-turns, stabilized by a conventional intraturn hydrogen bond. Considering the signs as well as magnitudes of the backbone torsion angles and the orientation of the central peptide bond, the overall mimicked topology resembles the type II beta-turn or type II Asx-turns. An analysis of Xaa-Thr sequences in high-resolution X-ray elucidated protein structures revealed the novel topology prevalence in functional proteins (unpublished). In view of indubitable structural as well as functional importance of nonconventional interactions in bioorganic and biomacromolecules, we intend to highlight the participation of Thr CgammaH in the creation of a short-range C=O . . . H--Cgamma -type interaction in peptides and proteins.     

Paternal signature in kin recognition cues of a social insect: concealed in juveniles,revealed in adults

Kin recognition is a key mechanism to direct social behaviours towards related individuals or avoid inbreeding depression. In insects, recognition is generally mediated by cuticular hydrocarbon (CHC) compounds, which are partly inherited from parents. However, in social insects, potential nepotistic conflicts between group members from different patrilines are predicted to select against the expression of patriline-specific signatures in CHC profiles. Whereas this key prediction in the evolution of insect signalling received empirical support in eusocial insects, it remains unclear whether it can be generalized beyond eusociality to less-derived forms of social life. Here, we addressed this issue by manipulating the number of fathers siring clutches tended by females of the European earwig, Forficula auricularia, analysing the CHC profiles of the resulting juvenile and adult offspring, and using discriminant analysis to estimate the information content of CHC with respect to the maternal and paternal origin of individuals. As predicted, if paternally inherited cues are concealed during family life, increases in mating number had no effect on information content of CHC profiles among earwig juveniles, but significantly decreased the one among adult offspring. We suggest that age-dependent expression of patriline-specific cues evolved to limit the risks of nepotism as family-living juveniles and favour sibling-mating avoidance as group-living adults. These results highlight the role of parental care and social life in the evolution of chemical communication and recognition cues.     

Trait‐mediated effects of predation across life‐history stages in container mosquitoes

1. It was determined if the predatory midge Corethrella appendiculata Grabham imposes a fitness cost in a native mosquito, Ochlerotatus triseriatus Say, and an invasive mosquito, Aedes albopictus Skuse. The hypothesis that decreased activity of immature prey in the presence of predator cues is associated with life history costs through all life cycle stages was tested. 2. In experiment 1, individual larvae of O. triseriatus or A. albopictus were raised in the presence or absence of predation cues at two resource levels. Prey were video recorded to detect behavioural responses and to measure development time, size at emergence, and adult longevity. In experiment 2, prey populations were reared in similar environments and the frequency of predator cue additions was varied. 3. Only O. triseriatus reduced its activity in the presence of predation cues. Predation cues were associated with longer immature development times and shorter adult life spans in O. triseriatus, whereas in A. albopictus, the cues were associated with a larger size of emerging adults. 4. In the present study, it was found that behavioural modifications during the larval stage can affect mosquitoes through multiple stages of their complex life cycle. The species‐specific behavioural differences are probably attributable to the longer evolutionary history O. triseriatus has with predators, relative to the invasive A. albopictus.     

A chemical signal of offspring quality affects maternal care in a social insect

Begging signals of offspring are condition-dependent cues that are usually predicted to display information about the short-term need (i.e. hunger) to which parents respond by allocating more food. However, recent models and experiments have revealed that parents, depending on the species and context, may respond to signals of quality (i.e. offspring reproductive value) rather than need. Despite the critical importance of this distinction for life history and conflict resolution theory, there is still limited knowledge of alternative functions of offspring signals. In this study, we investigated the communication between offspring and caring females of the common earwig, Forficula auricularia, hypothesizing that offspring chemical cues display information about nutritional condition to which females respond in terms of maternal food provisioning. Consistent with the prediction for a signal of quality we found that mothers exposed to chemical cues from well-fed nymphs foraged significantly more and allocated food to more nymphs compared with females exposed to solvent (control) or chemical cues from poorly fed nymphs. Chemical analysis revealed significant differences in the relative quantities of specific cuticular hydrocarbon compounds between treatments. To our knowledge, this study demonstrates for the first time that an offspring chemical signal reflects nutritional quality and influences maternal care.     

Behavioural ecology of defence in a risky environment: caterpillars versus ants in a Neotropical savanna

1. Predatory ants may reduce infestation by herbivorous insects, and slow‐moving Lepidopteran larvae are often vulnerable on foliage. We investigate whether caterpillars with morphological or behavioural defences have decreased risk of falling prey to ants, and if defence traits mediate host plant use in ant‐rich cerrado savanna. 2. Caterpillars were surveyed in four cerrado localities in southeast Brazil (70–460 km apart). The efficacy of caterpillar defensive traits against predation by two common ant species (Camponotus crassus, C. renggeri) was assessed through experimental trials using caterpillars of different species and captive ant colonies. 3. Although ant presence can reduce caterpillar infestation, the ants' predatory effects depend on caterpillar defence traits. Shelter construction and morphological defences can prevent ant attacks (primary defence), but once exposed or discovered by ants, caterpillars rely on their size and/or behaviour to survive (secondary defence). 4. Defence efficiency depends on ant identity: C. renggeri was more aggressive and lethal to caterpillars than C. crassus. Caterpillars without morphological defences or inside open shelters were found on plants with decreased ant numbers. No unsheltered caterpillar was found on plants with extrafloral nectaries (EFNs). Caterpillars using EFN‐bearing plants lived in closed shelters or presented morphological defences (hairs, spines), and were less frequently attacked by ants during trials. 5. The efficiency of defences against ants is thus crucial for caterpillar survival and determines host plant use by lepidopterans in cerrado. Our study highlights the effect of EFN‐mediated ant‐plant interactions on host plant use by insect herbivores, emphasizing the importance of a tritrophic viewpoint in risky environments.     

Maximum likelihood estimate of life expectancy in the prehistoric Jomon: Canine pulp volume reduction suggests a longer life expectancy than previously thought

Recent theoretical progress potentially refutes past claims that paleodemographic estimations are flawed by statistical problems, including age mimicry and sample bias due to differential preservation. The life expectancy at age 15 of the Jomon period prehistoric populace in Japan was initially estimated to have been ～16 years while a more recent analysis suggested 31.5 years. In this study, we provide alternative results based on a new methodology. The material comprises 234 mandibular canines from Jomon period skeletal remains and a reference sample of 363 mandibular canines of recent‐modern Japanese. Dental pulp reduction is used as the age‐indicator, which because of tooth durability is presumed to minimize the effect of differential preservation. Maximum likelihood estimation, which theoretically avoids age mimicry, was applied. Our methods also adjusted for the known pulp volume reduction rate among recent‐modern Japanese to provide a better fit for observations in the Jomon period sample. Without adjustment for the known rate in pulp volume reduction, estimates of Jomon life expectancy at age 15 were dubiously long. However, when the rate was adjusted, the estimate results in a value that falls within the range of modern hunter‐gatherers, with significantly better fit to the observations. The rate‐adjusted result of 32.2 years more likely represents the true life expectancy of the Jomon people at age 15, than the result without adjustment. Considering ～7% rate of antemortem loss of the mandibular canine observed in our Jomon period sample, actual life expectancy at age 15 may have been as high as ～35.3 years.     

The evolution of a complex trait: cuticular hydrocarbons in ants evolve independent from phylogenetic constraints

Cuticular hydrocarbons (CHCs) are ubiquitous and highly diverse in insects, serving as communication signal and waterproofing agent. Despite their vital function, the causes, mechanisms and constraints on CHC diversification are still poorly understood. Here, we investigated phylogenetic constraints on the evolution of CHC profiles, using a global data set of the species‐rich and chemically diverse ant genus Crematogaster. We decomposed CHC profiles into quantitative (relative abundances, chain length) and qualitative traits (presence/absence of CHC classes). A species‐level phylogeny was estimated using newly generated and previously published sequences from five nuclear markers. Moreover, we reconstructed a phylogeny for the chemically diverse Crematogaster levior species group using cytochrome oxidase I. Phylogenetic signal was measured for these traits on genus and clade level and within the chemically diverse C. levior group. For most quantitative CHC traits, phylogenetic signal was low and did not differ from random expectation. This was true on the level of genus, clade and species group, indicating that CHC traits are evolutionary labile. In contrast, the presence or absence of alkenes and alkadienes was highly conserved within the C. levior group. Hence, the presence or absence of biosynthetic pathways may be phylogenetically constrained, especially at lower taxonomic levels. Our study shows that CHC composition can evolve rapidly, allowing insects to quickly adapt their chemical profiles to external selection pressures, whereas the presence of biosynthetic pathways appears more constrained. However, our results stress the importance to consider the taxonomic level when investigating phylogenetic constraints.     

Socially cued anticipatory adjustment of female signalling effort in a moth

Juvenile population density has profound effects on subsequent adult development, morphology and reproductive investment. Yet, little is known about how the juvenile social environment affects adult investment into chemical sexual signalling. Male gumleaf skeletonizer moths, Uraba lugens, facultatively increase investment into antennae (pheromone receiving structures) when reared at low juvenile population densities, but whether there is comparable adjustment by females into pheromone investment is not known. We investigate how juvenile population density influences the ‘calling'' (pheromone-releasing) behaviour of females and the attractiveness of their pheromones. Female U. lugens adjust their calling behaviour in response to socio-sexual cues: adult females reared in high juvenile population densities called earlier and for longer than those from low juvenile densities. Juvenile density also affected female pheromonal attractiveness: Y-maze olfactometer assays revealed that males prefer pheromones produced by females reared at high juvenile densities. This strategic investment in calling behaviour by females, based on juvenile cues that anticipate the future socio-sexual environment, likely reflects a response to avoid mating failure through competition with neighbouring signallers.