蜜蜂蜂王信息素研究进展

综述了蜂王信息素的化学组成、特性、作用 ,以及与幼虫信息相互关系的研究进展 ,对蜂王信息素在养蜂和植物授粉中的应用做了介绍。西方蜜蜂的蜂王上颚腺信息素 (queen’smandibularglandpheromone,QMP)。QMP包含了 5种成分 :(E) -9-氧 -2 -癸烯酸 ( 9 ODA)、(R ,E) ( ) 和 (S,E) ( +) 9 羟基 -2 -癸烯酸 ( -9 HDA和 +9 HDA)、甲基p -羟基安息香酸盐 (HOB)、4-羟基 -3 -甲氧苯乙醇 (也称香草醇 ,HVA) ,9 ODA是其中最重要的成分 ;QMP的组分与蜜蜂的进化程度有关 ,进化程度越高则组分越复杂 ;QMP通过抑制保幼激素的产生来调节青年工蜂的个体发育。     

Sex pheromones of Callosobruchus subinnotatus and C. maculatus (Coleoptera: Bruchidae): congeneric responses and role of air movement

Females of Callosobruchus spp. are known to produce sex pheromones that attract males. These sex pheromones cannot be adopted for use in pest management without first investigating the responses of the males in the windless conditions of storage environments. Consequently, behavioural bioassays of Callosobruchus subinnotatus Pic males were conducted in an olfactometer in the absence of air-flow. Under these conditions males were found to be able to follow odour trails to the source. However, the latency period was longer in diffusional bioassays than for insects in a Y-tube olfactometer that provided directional wind cues. The highest percentage of males reached the pheromone source when components of the pheromones, (E)-3-methyl-2-heptenoic acid (E32A) and (Z)-3-methyl-2-heptenoic acid (Z32A), were formulated in a 50:50 or 25:75 ratio. Males of C. maculatus (Fabricius) responded to sex pheromone of C. subinnotatus, but males of C. subinnotatus did not respond to that of C. maculatus. The two sex pheromone components of C. subinnotatus are also constituents of C. maculatus sex pheromone. These two components may be potentially useful in monitoring the populations of both species in stored beans. It is postulated that (Z)-3-methyl-3-heptenoic acid (Z33A), the major component of the sex pheromone of C. maculatus, must have acted as an antagonist inhibiting response of C. subinnotatus to the sex pheromone of C. maculatus.     

Cracking the olfactory code of a butterfly: the scent of ageing

Although olfaction is a primary mode of communication, its importance in sexual selection remains understudied. Here, using the butterfly Bicyclus anynana, we address all the parameters of importance to sexual selection for a male olfactory signal. We show that variation in the male sex pheromone composition indicates male identity and male age. Courting males of different ages display small absolute (c. 200 ng) but large relative (100%) change of one specific pheromone component (hexadecanal) which, unlike the other components, showed no heritability. Females prefer to mate with mid-aged over younger males and the pheromone composition is sufficient to determine this preference. Surprisingly refined information is thus present in the male olfactory signal and is used for sexual selection. Our data also reveal that there may be no 'lek paradox' to resolve once the precise signal of importance to females is identified, as hexadecanal is, as expected, depleted in additive genetic variation.     

Pheromonal composition of two species of african Amblyomma ticks: similarities,differences and possible species specific components

Two species of bont ticks, Amblyomma hebraeum and Amblyomma variegatum, common to Africa were studied to compare types and quantities of compounds known or believed to serve as components of the attraction-aggregation-attachment pheromone (AAAP). A complex of attraction, aggregation and attachment stimulating pheromone components are used by these to detect hosts, mates and, perhaps, minimize interspecific breeding. Solvent extraction of pheromone emitting ticks followed by gas chromatography and mass spectrometry revealed little qualitative difference in the composition of the AAAP in these two species. However, subtle differences in the relative proportions of the pheromonal blend are noted suggesting that such differences in concentration may facilitate species-specific discrimination during aggregation and attachment. Differences in the relative abundance of benzaldchyde and methyl salicylate in the males of the two species were especially noteworthy. Possible means by which such differences in phenolic compound composition may affect the behavior of these ticks are discussed.     

Pheromone production, male abundance, body size, and the evolution of elaborate antennae in moths

The males of some species of moths possess elaborate feathery antennae. It is widely assumed that these striking morphological features have evolved through selection for males with greater sensitivity to the female sex pheromone, which is typically released in minute quantities. Accordingly, females of species in which males have elaborate (i.e., pectinate, bipectinate, or quadripectinate) antennae should produce the smallest quantities of pheromone. Alternatively, antennal morphology may be associated with the chemical properties of the pheromone components, with elaborate antennae being associated with pheromones that diffuse more quickly (i.e., have lower molecular weights). Finally, antennal morphology may reflect population structure, with low population abundance selecting for higher sensitivity and hence more elaborate antennae. We conducted a phylogenetic comparative analysis to test these explanations using pheromone chemical data and trapping data for 152 moth species. Elaborate antennae are associated with larger body size (longer forewing length), which suggests a biological cost that smaller moth species cannot bear. Body size is also positively correlated with pheromone titre and negatively correlated with population abundance (estimated by male abundance). Removing the effects of body size revealed no association between the shape of antennae and either pheromone titre, male abundance, or mean molecular weight of the pheromone components. However, among species with elaborate antennae, longer antennae were typically associated with lower male abundances and pheromone compounds with lower molecular weight, suggesting that male distribution and a more rapidly diffusing female sex pheromone may influence the size but not the general shape of male antennae.     

Movement of honey bee (Apis mellifera L.) queen mandibular gland pheromone in populous and unpopulous colonies

The mode of intranest transfer of the honey bee queen mandibular gland pheromone complex (QMP) was investigated in unpopulous and populous, slightly congested colonies, using synthetic QMP containing tritiated 9-keto-2(E)-decenoic acid, one of the QMP components. Radiolabel was rapidly transported from the center to the peripheral regions of the nest, and in a manner consistent with worker to worker transport. Population size and congestion had no effect on the relative rates of movement from the center to the periphery of the nest or on the mean amounts of radiolabel on individual bees. However, a significantly smaller proportion of the workers in the populous colonies received detectable amounts of radiolabel than in the uncongested colonies, and workers carrying especially large amounts of radiolabel were less numerous in the crowded colonies. It is suggested that, at the stage of colony development that the colonies were in, population size has more of an effect on intranest pheromone transmission than does crowding. Interference with pheromone transfer may occur only at higher levels of congestion than were created, and nearer to the reproductive phase of colony development. An alternative hypothesis is that colony crowding does not significantly affect QMP transport and that the onset of reproductive queen rearing may be associated more with changes in worker thresholds of response to QMP.     

The mode of pheromone evolution: evidence from bark beetles

Sex and aggregation pheromones consist of species-specific blends of chemicals. The way in which different species' blends have evolved has been the subject of some debate. Theoretical predictions suggest that differences between species have arisen not through the accruing of small changes, but through major shifts in chemical composition. Using data on the aggregation pheromones of 34 species of bark beetle from two genera, Dendroctonus and Ips, we investigated how the distributions of the chemical components of their pheromone blends mirror their phylogenetic relationships. We tested whether there were consistent patterns that could be used to help elucidate the mode of pheromone evolution. Although there were obvious differences in pheromone blends between the two genera, the differences between species within each genus followed a less clear phylogenetic pattern. In both genera, closely related species are just as different as more distantly related species. Within Dendroctonus, particularly, most chemical components were distributed randomly across the phylogeny. Indeed, for some chemicals, closely related species may actually be more different than would be expected from a random distribution of chemical components. This argues strongly against the idea of minor shifts in pheromone evolution. Instead, we suggest that, within certain phylogenetic constraints, pheromone evolution in bark beetles is characterized by large saltational shifts, resulting in sibling species being substantially phenotypically (i.e. pheromonally) different from one another, thus agreeing with theoretical predictions.     

PHEROMONES AND CHEMICAL ECOLOGY OF LOCUSTS

Modern studies of chemical ecology and behaviour of the locusts Schistocerca gregaria and Locusta migratoria in the laboratory need to be more closely coupled with field experiments and observations. The life history relating to oviposition, transformation to gregarious phases, and adult maturation mediated by pheromones is reviewed. The principles of pheromone isolation and identification are discussed. The long-term effects of the gregarization pheromone on the physiology are presented, with discussion of morphological changes, chiasma frequency increases, and synchronization of moulting induced by the pheromone. Isolation of the purported gregarization pheromone, locustol, from faeces is discussed in regard to inconsistent effects. Other more immediate effects of the pheromone on the social (gregarious) behaviour and the isolation of possible pheromone components different from but related to locustol are presented. It is stressed that more rigorous isolation studies should be undertaken to resolve conflicting reports and methodological problems. The possibility of an anti-gregarization pheromone or solitarizing pheromone is discounted. The source and biosynthesis of locustol (or gregarization pheromone) from degradation of lignin by symbiotic bacteria is discussed. Theories of reception of the gregarization pheromone such as inhalation through the spiracles or sensory perception by the antennae are presented. Also an internal mechanism involving cAMP and/or corpora allata may be induced by gregarization pheromone to effect the physiological phase changes. The advantages to an individual of reception of the gregarization pheromone from a group of gregarious and pre-migrating locusts is discussed. Also the possible benefits of gregarious behaviour, phase polymorphism and migration are dealt with. An adult (sexual) maturation pheromone has long-term effects on reducing the period of maturation, and immediate effects on the behavioural vibration response. The epidermal source of the pheromone and glandular cells responsible for the production of the pheromone are discussed. The reception and internal mechanisms of response via the corpora allata are mentioned. The benefits to individuals of synchronized and rapid adult maturation in a gregarious group are considered. An oviposition-stimulating pheromone produced by the male accessory reproductive glands appears to be a proteinaceous substance of large molecular weight. On the other hand, an oviposition-aggregating pheromone volatilizes from epidermal areas of either sex and causes higher oviposition rates in the area of release. The behavioural and ecological aspects of this pheromone are discussed. Several other possible pheromones and semiochemicals are discussed, such as a long-range sex pheromone, sex-recognition pheromone, grass odours and feeding stimulants and deterrents. Several possible control strategies using locust pheromones are considered. The general conclusion is that the chemical isolation of the various pheromones is necessary before further progress can be achieved on the source and biosynthesis of pheromone, reception of pheromone, behavioural effects of pheromone, and control measures.     

Characterization of the general odorant-binding protein 2 in the molecular coding of odorants in Mamestra brassicae.

The general odorant-binding protein 2 of Mamestra brassicae males has been purified from antennal extracts and examined in binding assays with pheromone components of this species and a behavioral antagonist, cis-11-hexadecenol. The protein showed high affinity for the latter compound and no affinity for the pheromone components. In addition, expression of the protein, studied by in situ hybridization, was restricted to the long sensilla trichodea, which house the neuron that responds to cis-11-hexadecenol. The expression in a functionally defined population of sensilla, together with binding specificity and previous electrophysiological data, suggest an unsuspected role for the general odorant-binding protein 2 in M. brassicae. It may be involved in the transduction process for the behavioral antagonist to which neurons are specifically tuned and always cocompartmentalized in long trichodeal hairs, with neurons responding to the major pheromonal compound, cis-11-hexadecenyl acetate. These data are consistent with the involvement of odorant-binding proteins in the fine discrimination between pheromone and antagonist, which is related to avoidance of interspecific mating mistakes.     

Artificial night lighting disrupts sex pheromone in a noctuid moth

1. One major, yet poorly studied, change in the environment is the increase in nocturnal light pollution. Although this strongly alters the habitat of nocturnal species, the ecological consequences are poorly known. Moths are well known to be attracted to artificial light sources, but artificial light may affect them in other ways as well. 2. In this study, female Mamestra brassicae moths were subjected to various types of low‐intensity artificial night lighting with contrasting spectral compositions (green‐rich, red‐rich, warm white) or to a dark control treatment and the effects on their sex pheromone production and composition were tested. 3. Artificial night lighting reduced sex pheromone production and altered the chemical composition of the pheromone blend, irrespective of spectral composition. Specifically, amounts of the main pheromone component Z11‐16:Ac were reduced, while the deterring compounds Z9‐14:Ac, Z9‐16:Ac, and Z11‐16:OH were increased relative to Z11‐16:Ac when females were kept under artificial light. These changes may reduce the effectiveness of the sex pheromones, becoming less attractive for males. 4. These results show for the first time that artificial light at night affects processes that are involved in moth reproduction. The potential for mitigation through manipulation of the spectral composition of artificial light appears limited.     

Context‐dependent alarm signalling in an insect

Animals often respond to danger by raising alarm to inform others. Alarm signals come in many different forms, such as visual or mechanical display, sound or odour. Some animals produce vocal alarm signals that vary with the level of danger. For chemical alarm signals, virtually nothing is known about such context‐dependent signalling due to a general notion that alarm pheromones have fixed compositions. Here, we show that larvae of the Western Flower Thrips (Frankliniella occidentalis) produce an alarm pheromone whose composition varies with the level of danger they face: the presence of a relatively harmless predator or a very dangerous predator, that is either actually attacking or not. The frequency of alarm pheromone excretion increases with the level of danger. Moreover, the composition of excreted alarm pheromone varies in the relationship between total and relative amount of the putative two components, decyl acetate (DAc) and dodecyl acetate (DDAc). When pheromone is excreted with a predator present but not attacking, the percentage DDAc increases with the total amount of pheromone. When a predator does attack, however, the relationship between percentage DDAc and total amount of pheromone is reversed. Taken together, the alarm signal of thrips larvae appears to be context dependent, which to our knowledge is the first report of context‐dependent composition of an alarm pheromone.     

Geographical variation in female sex pheromones of the rice leaffolder moth, Cnaphalocrocis medinalis: identification of pheromone components in Japan

Sex pheromone components of the Japanese rice leaffolder moth, Cnaphalocrocis medinalis Guenée (Lepidoptera: Pyralidae) were identified from ovipositor extracts of virgin females as (Z)-11-octadecenal, (Z)-13-octadecenal, (Z)-11-octadecen-1-ol and (Z)-13-octadecen-1-ol at a ratio of 11:100:24:36 by GC-EAD, GC, GC-MS. The total amount was estimated to be ca.0.9 ng/female. Field bioassays in Kagoshima, Japan, showed that the two aldehydes are essential for male attraction and the alcohols may have a synergistic effect on the aldehydes. A rubber septum containing 0.9 mg of the four components at the natural ratio was shown to be an effective lure for monitoring this pest in Japan. The above four components are quite different from the sex pheromone components reported previously for the same species of either Philippine or Indian origin; components were shown to be (Z)-11-hexadecenyl acetate and (Z)-13-octadecenyl acetate at a ratio of 98:2 in the Philippine blend and 1:10 in the Indian blend. Furthermore, in the field tests in Japan, neither the Philippine blend nor the Indian blend showed any attractive activity, while the Japanese blend attracted significant numbers of male moths. These results suggest that there are remarkable geographical variations in the sex pheromone composition of this species or there are several distinct species using different sex pheromone blends.     

The chiasma-inducing pheromone of locusts

To the two quantitative criteria for phase transformation of locusts, viz. nymphal colour and adult morphometric ratios, the change in chiasma frequencies during meiosis in the male can now be added: gregarization or swarming induces an increase in chiasma formation. The atmosphere around crowded locusts contains a pheromone which is absorbed and causes a reactive haemolymph in locust hoppers. This haemolymph may be injected into solitarious hoppers or nymphs to induce increased chiasma formation. Certain solvents like risella oil and dimethyl sulphoxide will extract the pheromone from such an atmosphere, and bio-assays with these extracts have been demonstrated to be chiasma-inducing. Such extracts are, however, not chiasma-inducing in the albino mutant, but the haemolymph from crowded normal-coloured hoppers is reactive in the albino hopper. In addition the albino mutant is solitaria-like in both morphometric ratios and chiasma frequencies. These data indicate a relationship between the melanization cycle and chiasma induction: the previously postulated melanin-inducing pheromone and this chiasma-inducing pheromone are probably identical. On the other hand, this pheromone of nymphs and the sex-maturation pheromone of adults are not identical, although the latter may be a modified form of the other. The effect of the nymphal pheromone on adult morphometric ratios is not clear although some correlation has been shown to exist between raised chiasma frequencies in treated solitaries and advances towards gregaria ratios. It may be that in this pheromone we have, what may be called, a gregarizing principle.     

Isolation and identification of terpenoid sex pheromone components from extracts of hemolymph of males of the Caribbean fruit fly

Extracts obtained from hemolymph of sexually mature males of the Caribbean fruit fly Anastrepha suspensa contained four biologically important terpenoid components of the sex pheromone. The four components were identified as farnesene, bisabolene, anastrephin, and epianastrepin based on their relative retention indexes from capillary gas chromatography analysis, using both apolar and polar phase columns and their chemical ionization (isobutane) mass spectra. The ratio of the components in extracts of hemolymph was the same as the ratio present in the volatile blend of pheromone released by sexually mature males during the reproductive period. Studies conducted to determine the effect of age on amounts of these components in hemolymph indicated that they increased from undetectable levels on the day of adult emergence to maximum levels on day eight. The increases in amounts of the components present in hemolymph with increasing age were correlated with increases in amounts of volatile pheromone released by males. Time of day studies showed that the amounts of these components in hemolymph followed the daily pattern of release of volatile pheromone components. Other components of the sex pheromone including ocimene, (Z)-3-nonen-1-ol, (Z,Z)-3,6-nonadien-1-ol and suspensolide were not found in extracts of hemolymph. The data suggest that the hemolymph plays a role in the transport of these pheromone components during sexual signalling. Arch. Insect Biochem. Physiol. 42:225-232, 1999. Published 1999 Wiley-Liss, Inc.     

THE EVOLUTION OF SOCIAL SIGNALS: MORPHOLOGICAL,FUNCTIONAL, AND GENETIC INTEGRATION OF THE SEX PHEROMONE IN NAUPHOETA CINEREA

Social signals that mediate intraspecific interactions can be complex, conveying considerable information concerning the probable behavior of individuals and minimizing overt aggression and wasted energy. In the cockroach Nauphoeta cinerea, male-male competition and female mate choice are mediated by a multicomponent male-produced sex pheromone. In this study, I examine variation in this pheromone. First I measure differences among males in both individual pheromone compounds and the overall composition of the pheromone. Principal component analysis is used to quantify and describe pheromone composition. Next, I explore some of the causes and consequences of this variation by examining the pheromone of males with different social experiences. Compared to subordinate males, dominant males have significantly less variable quantities of the individual pheromone compounds and are significantly less variable in the composition of their pheromone. Because of an association between status and mating success, male-male competition can result in stabilizing sexual selection on the sex pheromone. Finally, I test the hypothesis that the pheromone compounds evolve in a manner consistent with their function. As predicted for morphologically integrated characters, the patterns of phenotypic, genetic, and environmental correlations among my measures of pheromone compounds and composition match functional patterns suggested by this study and the developmental patterns demonstrated in my previous studies. Based on these studies of the N. cinerea sex pheromone, I argue that stabilizing sexual selection shapes the evolution of pheromonal communication involved in social interactions among male N. cinerea. Further, I argue that coordinated evolution of social signals may be possible due to the morphological integration of their multiple compounds.     

Peripheral Mechanisms of Pheromone Reception in Moths

Moths pheromones mostly consist of two or a few chemical componentsin a species-specific ratio. Each component is perceived bya particular type of receptor cell. Some pheromone componentscan inhibit the behavioral responses to other pheromone components.A single pheromone molecule is sufficient to elicit a nerveimpulse. The dose-response curve of single pheromone receptorneurons increases over many decades of stimulus intensity. Pheromonereceptor cells can resolve single stimulus pulses up to a frequencyof 10 pulses/s. Electrophysiological and biochemical studieson perireceptor events suggest that the pheromone moleculesinteract with the receptor cell while bound to a reduced formof the pheromone binding protein. The enzymatic degradationof pheromone found on the antennae is much too slow to accountfor the decline of the receptor potential after end of stimulation.The postulated rapid deactivation of the odor molecules adsorbedmight be performed by an oxidation of the pheromone bindingprotein. Several second messenger systems seem to be involvedin the cellular transduction mechanism (IP₃ diacylglycerol,cGMP, Ca₂₊). It is, however, not excluded that pheromone moleculescan gate single ion channels directly and thus elicit the elementaryreceptor potentials, observed at weak stimulus intensities.Chem. Senses 21: 257–268, 1996.     

Chemical constraints on the evolution of olfactory communication channels of moths

It is estimated there are over 100,000 moth species (Lepidoptera) that produce sex pheromones comprising communication channels used in specific-mate-recognition systems (SMRS) involved in pre-zygotic mating isolation and speciation. About 1572 moth species have been found to use 377 pheromone components, the majority being alcohols, aldehydes or acetate esters of olefinic chains of 10-22 carbons. Since there are limited numbers of possible unsaturated (double bonded) E- and Z-configured isomers of these chains, there may be constraints on incipient species evolving new pheromone components, especially if they are utilized by existing species. Here I develop algorithms that count and name all possible structures of chains with or without a functional group. The results show that for acetate esters there are only seven or nine monounsaturated isomers of six or seven carbon chains, respectively, suggesting use of these compounds could limit the number of communication channels available for radiation of new species (no moths use these short chains). For commonly used 14-carbon chains with an acetate ester functional group, and 1-3 unsaturations, there are 1039 isomers. A total of 2,096,883 isomers were counted from all multiply unsaturated 10-22-carbon chain acetate esters. The number of possible signals quickly extends into millions when considering pheromone blends of 2-4 components used by most species. There should be little chemical constraint on evolution of new communication channels based on compounds of 10 or more carbons, even for closely related species (e.g. ermine moths, Yponomeuta).