Asymmetrical generalisation between pheromonal and floral odours in appetitive olfactory conditioning of the honey bee (Apis mellifera L.)

The capacity to generalise between similar but not identical olfactory stimuli is crucial for honey bees, allowing them to find rewarding food sources with varying volatile emissions. We studied bees' generalisation behaviour with odours having different biological values: typical floral odours or alarm compounds. Bees' behavioural and peripheral electrophysiological responses were investigated using a combined proboscis extension response conditioning-electroantennogram assay. Bees were conditioned to pure linalool (floral) or to pure isoamyl acetate (alarm) and were tested with different concentrations of both compounds. Electrophysiological responses were not influenced by conditioning, suggesting that the learning of individual compounds does not rely on modulations of peripheral sensitivity. Behaviourally, generalisation responses of bees conditioned to the alarm compound were much higher than those of bees conditioned to the floral odour. We further demonstrated such asymmetrical generalisation between alarm and floral odours by using differential conditioning procedures. Conditioning to alarm compounds (isoamyl acetate or 2-heptanone) consistently induced more generalisation than conditioning to floral compounds (linalool or phenylacetaldehyde). Interestingly, generalisation between the two alarm compounds, which are otherwise chemically different, was extremely high. These results are discussed in relation to the neural representation of compounds with different biological significance for bees.     

Genetic and developmental factors in the olfactory response of Drosophila melanogaster larvae to alcohols.

Olfactory responses of Drosophila melanogaster larvae to a homologous series of primary alcohols (methanol ... decanol) were tested. Alcohols at either extreme of the chain lengths studied (methanol, ethanol and decanol) evoked no significant responses. Heptanol and nonanol both produced dose-independent responses, larvae being attracted to heptanol and repulsed by nonanol. The remaining alcohols elicited dose-related attractive responses. Responses to hexanol and nonanol decline with increasing larval age. Genetic differences were found for the response to heptanol, with larvae from a Japanese strain, Katsunuma, being indifferent to this substance. Chromosome exchange revealed that a major factor involved in the response to heptanol is located on chromosome II; factors on chromosome III quantitatively modulate this response. Three mutant strains were isolated following EMS mutagenesis of chromosome III. These three strains, IndifferentA, IndifferentB and IndifferentC, show incomplete or total anosmia when stimulated with nonanol. Adult flies from these strains show similar effects. IndifferenB and C strains are dominant over the Canton-S control strain; the IndifferentA strain shows semi-dominance. Results are discussed in the light of the ecology of Drosophila larvae and the relation between olfactory stimulus and receptor conformation and number.     

麦麸及其挥发性物质对二点委夜蛾幼虫的引诱作用

【目的】二点委夜蛾Athetis lepigone是我国夏玉米上的一种新害虫,对玉米苗危害严重。由于其幼虫常隐蔽在收割机收获小麦后留在田间的麦秸和麦糠下,防治非常困难,因此开展二点委夜蛾幼虫引诱剂的研究具有重要的意义。【方法】本实验用“Y”型嗅觉仪测定了二点委夜蛾幼虫对麦秆﹑麦糠﹑玉米苗﹑麦麸﹑炒麦麸5种试材的行为反应;利用顶空固相微萃取(HS-SPME)与气相色谱 质谱联用仪（GC-MS）分析,比较了炒麦麸、麦麸中挥发物的成分,并从中挑选出6种化合物单体、8种混合物进行室内诱集实验。【结果】5种试材中炒麦麸引诱效果最好。6种单体中, 2-庚酮在5 μg/μL浓度下的引诱效果最好,引诱率为75.9%。8种混合物中,2 庚酮+苯甲醛+对二甲苯的引诱效果较好,引诱率为72.7%,但引诱效果不如2 庚酮单体。【结论】结果说明,2-庚酮对二点委夜蛾幼虫有强烈的引诱作用,从而为二点委夜蛾幼虫的田间诱杀技术提供了研究基础。     

The significance of background odour for an egg parasitoid to detect plants with host eggs

Scots pine has been shown to produce a volatile bouquet that attracts egg parasitoids in response to oviposition of the herbivorous sawfly Diprion pini. Previous analyses of headspace volatiles of oviposition-induced pine twigs revealed only quantitative changes; in particular, the sesquiterpene (E)-beta-farnesene was emitted in significantly higher quantities by oviposition-induced pine. Here we investigated whether (E)-beta-farnesene attracted the egg parasitoid Chrysonotomyia ruforum. We tested the behavioural response of C. ruforum females to different concentrations of (E)-beta-farnesene. Egg parasitoids did not respond to this sesquiterpene at either concentration tested. However, they did respond significantly to (E)-beta-farnesene when this compound was offered in combination with the volatile blend emitted from pine twigs without eggs. This response was dependent on the applied concentration of (E)-beta-farnesene. Further bioassays with other components [(E)-beta-caryophyllene, delta-cadinene] of the odour blend of pine were conducted in combination with the volatile blend from egg-free pine as background odour. None of the compounds tested against the background of odour from an egg-free pine twig were attractive to the egg parasitoid. These results suggest that the egg parasitoids responded specifically to (E)-beta-farnesene, but only when this compound was experienced in the 'right' context, i.e. when contrasted with a background odour of non-oviposition-induced pine volatiles.     

Oriented responses of grapevine moth larvae Lobesia botrana to volatiles from host plants and an artificial diet on a locomotion compensator

Larvae of the grapevine moth Lobesia botrana (Lepidoptera: Tortricidae) are a major pest of vine, Vitis vinifera. As larvae have limited energy reserves and are in danger of desiccation and predation an efficient response to plant volatiles that would guide them to food and shelter could be expected. The responses of starved 2nd or 3rd instar larvae to volatile emissions from their artificial diet and to single host plant volatiles were recorded on a locomotion compensator. Test products were added to an air stream passing over the 30 cm diameter servosphere. The larvae showed non-directed walks of low rectitude in the air stream alone but changed to goal-oriented upwind displacement characterised by relatively straight tracks when the odour of the artificial diet and vapours of methyl salicylate, 1-hexanol, (Z)-3-hexen-1-ol, terpinen-4-ol, 1-octen-3-ol, (E)-4,8-dimethyl-1,3,7-nonatriene and (Z)-3-hexenyl acetate were added to the air stream. This chemoanemotactic targeted displacement illustrates appetence for certain volatile cues from food by starved Lobesia larvae. Analysis of the larval behaviour indicates dose dependent responses to some of the host plant volatiles tested with a response to methyl salicylate already visible at 1 ng, the lowest source dose tested. These behavioural responses show that Lobesia larvae can efficiently locate mixtures of volatile products released by food sources as well as single volatile constituents of their host plants. Such goal-oriented responses with shorter travel time and reduced energy loss are probably of importance for larval survival as it decreases the time they are exposed to biotic and abiotic hazards.     

Oriented responses of carrot fly larvae, Psila rosae, to plant odours, carbon dioxide and carrot root volatiles

ABSTRACT. The responses of carrot fly larvae, Psila rosae , to host plant volatiles were studied. Two criteria for an oriented response were used: one based on whether an individual reached within 1.5 cm of the odour source and one involving analysis of the track patterns of larvae within 3 cm of the odour source. All larval instars were attracted to both host and non-host plant material, though to a lesser degree by the latter; they were also attracted to CO₂, but not to N₂. A steam volatile fraction of carrot roots, and methyl eugenol, a volatile component of this fraction, also attracted the larvae, differentially at different concentrations. The possible roles of both CO₂ and the host-specific secondary plant volatiles in the host plant finding behaviour of soil inhabiting arthropods are discussed.     

Olfactory coding in a simple system: adaptation in Drosophila larvae.

Drosophila melanogaster larvae were pre-stimulated with high concentrations of six homologous alcohols (C4-C9) and then tested for adaptation and cross-adaptation using these same alcohols, four aliphatic n-acetates and three acids. Pre-stimulation with hexanol effectively reduced to zero (abolished) test responses to all six alcohols, whereas test responses to hexanol were only affected by pre-stimulation with hexanol. This substance appears to play a fundamental role in the organization of the larval olfactory system. Test responses to butanol and pentanol, and the effect of pre-stimulation with butanol and pentanol, were not significantly different, indicating that they are sensory equivalents. Heptanol, octanol and nonanol induce a complex set of responses among one another. Cross-adaptation between functional groups was observed, in particular following pre-stimulation with hexanol, but there was also evidence that functional groups are coded separately. A model of olfactory processing in the fruitfly maggot is presented that explains the data and provides predictions for future anatomical, genetic and electrophysiological studies.     

Feel the heat: The effect of temperature on development, behavior and central pattern generation in 3rd instar Calliphora vicina larvae

Like in all poikilothermic animals, higher temperatures increase developmental rate and activity in Calliphora vicina larvae. We therefore could expect temperature to have a persistent effect on the output of the feeding and crawling central pattern generators (CPGs). When confronted with a steep temperature gradient, larvae show evasive behavior after touching the substrate with the cephalic sense organs. Beside this reflex behavior the terminal- and dorsal organ might also mediate long term CPG modulation. Both organs were thermally stimulated while their response was recorded from the maxillary- or antennal nerve. The terminal organ showed a tonic response characteristic while the dorsal organ was not sensitive to temperature. Thermal stimulation of the terminal organ did not affect the ongoing patterns of fictive feeding or crawling, recorded from the antennal- or abdominal nerve respectively. A selective increase of the central nervous system (CNS) temperature accelerated the motor patterns of both feeding and crawling. We propose that temperature affects centrally generated behavior via two pathways: short term changes like thermotaxis are mediated by the terminal organ, while long term adaptations like increased feeding rate are caused by temperature sensitive neurons in the CNS which were recently shown to exist in Drosophila larvae.     

葡萄酒相关酵母的香气形成及香气特征

一系列相关微生物的代谢及相互作用贯穿在葡萄酒的酿造过程中,其中各种酵母菌的代谢产物是影响葡萄酒感官特征的关键,不同菌种代谢产生不同的挥发性物质,造成对葡萄酒香气的最直接影响。介绍了酿酒酵母以及非酿酒酵母的代谢特征与差别,由此所引起的葡萄酒气味的不同表现,以及发酵过程中不同菌种的相互作用,总结了近些年来对非酿酒酵母酿酒特性的研究与利用。目前已证实非酿酒酵母的一些代谢特征对葡萄酒香气和香气结构具有一些积极的作用,并且酿酒酵母和有些非酿酒酵母的结合使用对改善葡萄酒感官也具有良好的作用与发展前景。     

葡萄酒相关酵母的香气形成及香气特征

一系列相关微生物的代谢及相互作用贯穿在葡萄酒的酿造过程中,其中各种酵母菌的代谢产物是影响葡萄酒感官特征的关键,不同菌种代谢产生不同的挥发性物质,造成对葡萄酒香气的最直接影响。介绍了酿酒酵母以及非酿酒酵母的代谢特征与差别,由此所引起的葡萄酒气味的不同表现,以及发酵过程中不同菌种的相互作用,总结了近些年来对非酿酒酵母酿酒特性的研究与利用。目前已证实非酿酒酵母的一些代谢特征对葡萄酒香气和香气结构具有一些积极的作用,并且酿酒酵母和有些非酿酒酵母的结合使用对改善葡萄酒感官也具有良好的作用与发展前景。     

The attractiveness of different odour sources from the fruit–host complex on Leptopilina boulardi, a larval parasitoid of frugivorous Drosophila spp.

In parasitoid insects, successful offspring development depends on the female’s ability to find a suitable host. Specific recognition is often based on responses to olfactory cues, but their source and nature have rarely been determined. –This paper deals with the recognition of odours involved in host location by Leptopilina boulardi[Barbotin, Carton & Kelner-Pillault] (Hymenoptera: Eucoilidae), a larval parasitoid of Drosophila species that develops in mature fruits. The nature and origin of volatile stimuli recognized among odours of the host–fruit complex, and the effect of learning on this recognition, were investigated. Oriented responses to these odours were observed in a four-armed olfactometer and were analysed with the observer software (Noldus Information Technology). Fruit odours alone (banana and pear) were not spontaneously attractive to naive parasitoids, whereas naturally-infested bananas were highly attractive. The attraction was related to the odour that adult Drosophila left on the substrate but not to Drosophila oviposition activity or larval development. A synergism between some fruit odours (banana and pear) and the odour left by adult Drosophila on damp filter paper was observed. However, when testing a non-fruit substrate (mushroom), no synergism was observed. Thus, female L. boulardi may innately recognize host–food substrate odours associated with odours from the adult stage of their host. In addition, an oviposition experience on an infested banana allows L. boulardi females to memorise the fruit odour itself through associative learning. The adaptive significance of this process is discussed.     

Physico chemical alterations in desugared buffalo milk after fermentation withSaccharomyces fragilis

Summary Buffalo milk was completely desugared by fermentation for 17 h withSaccharomyces fragilis. A thin-layer chromatographic profile indicated the total absence of lactose, galactose, glucose and any oligosaccharides. Desugaring by fermentation did not alter the lipid composition or the protein content of the milk. Gas chromatographic studies indicated the formation of volatile compounds such as acetaldehyde, ethyl acetate, ethyl alcohol, n-propanol, n-butanol and isoamyl alcohol. A pleasant fruity odour observed in the initial stage of fermentation changed to an overripe fruity odour on fermentation. The volatile compounds formed disappeared on spraydrying the milk, leaving only a residual yeasty odour in the desugared milk powder.     

The larval crustacean dorsal organ and its relationship to the trilobite median tubercle

In the dorsal median anterior region of the head, the larvae of decapod Crustacea possess a discrete organ: a glandular-sensory complex which is not present in post-larval juvenile forms. Its external topography reveals the presence of one or two central pores (Brachyura, Anomura) or a poreless central area (Macrura) surrounded by four equidistant plate-pits which contain a central pimple or 'nipple'. The Macrura plate-pits lack the central pimple but develop a row of diminutive pegs. No definite function can yet be ascribed to the organ. The dorsal organ described here in Decapoda larvae closely resembles the median sensory tubercle found on the cephalon of some trilobites, especially Trinucleidae. In view of the possible homology between these structures in the two groups, previous interpretation of the trilobite organ as a median eye tubercle must be rejected.     

Odour representation in honeybee olfactory glomeruli shows slow temporal dynamics: an optical recording study using a voltage-sensitive dye

Stimulation with odours has been shown to elicit characteristic patterns of activated glomeruli in the antennal lobe (AL) of honeybees. In this study we show that these patterns are dynamic in a time window of 2-3 s after stimulus onset. We measured changes in the averaged membrane potential of all cells in the glomerular neuropil by optical imaging of the voltage-sensitive dye RH795 using a slow scan CCD camera (3 frames/s). The four substances 1-hexanol, hexanal, citral and clove-oil as well as the binary mixtures hexanol+hexanal and hexanol+citral were used as stimuli (2 s stimulus duration). We found that: (1) every odour elicited an odour-specific activity pattern, and conversely every glomerulus had a characteristic odour response profile; (2) some glomeruli had a tonic, some a phasic-tonic, and some a slow phasic response pattern; (3) the difference between the glomerular response patterns increased within 2 s of stimulus presentation, which suggests that odour representations became more characteristic over stimulus time; and (4) the responses to odorant mixtures were complex and glomerulus-dependent: some responses correspond to the sum of the compounds' responses, some to the response of one of the components.     

Variability and genetic components of innate fruit odour recognition in a parasitoid of Drosophila

Abstract This study aims to investigate the genetic variability of olfactory responses to odours from the host habitat, involved in host selection by insect parasitoids. The probing response of females to fruit and non-fruit odours was studied in Leptopilina boulardi Barbotin et al . (Hymenoptera, Figitidae), a parasitoid of the frugivorous Drosophila melanogaster larvae. Responses of two strains (from Nasrallah, Tunisia, and from Brazzaville, Congo) and their hybrids were compared. Probing is a searching mode by repeated ovipositor insertions into the breeding substrate to locate digging larvae. Most Brazzaville females exhibited innate ovipositor probing into agar in response to the fruit aromas (banana, orange, pear, prickly pear), whereas this was rare in the Nasrallah strain. These four fruits are all breeding sites for D. melanogaster, whereas mushroom is not. Mushroom odour did not trigger any probing response in either strain. In addition, the latency of the probing response was shorter in Brazzaville females. Results from hybrid females (F1 and Backcross) showed that both probability to probe in response to a fruit aroma and latency of the response were strongly heritable. Regarding the probability to probe, there was an apparent complete dominance of Nasrallah characteristics, whereas additive inheritance was found in the Brazzaville lineage. The contrast between reciprocal F1 hybrids suggested a strong interaction with non-chromosomal factors. It is suggested that a cytoplasmic factor in the Nasrallah strain decreases females' propensity to probe in response to significant odours. The latency of probing appeared to be under a chromosomal influence. Thus, this study identifies a strong genetic component in the responses to odours from the host habitat. It also provides information on the genetic basis of the two characters studied.     

A sensory map based on velocity threshold of sensory neurones from a chordotonal organ in the tailfan of the crayfish

The central projections of sensory neurones innervating a strand chordotonal organ (CO) in the tailfan of the crayfish, Procambarus clarkii (Girard) have been investigated. The CO monitors movement of the exopodite of the tailfan relative to the endopodite. Intracellular recording and staining were used to characterise the response of the sensory neurones to applied stretches of the chordotonal organ and to reveal their morphology. Two gross morphological types of afferents were found: those that terminated in the terminal (6th) abdominal ganglion on the side ipsilateral to the sensory receptor, and those that had branches in the terminal ganglion and an intersegmental axon that ascended rostrally. Afferents responded to position, velocity and direction of imposed CO displacement. Afferents with particular physiological properties had similar morphologies in different crayfish. Irrespective of their directional responses, afferents had central projection areas dependent upon their velocity thresholds. Many afferents responded only during movement of the CO, and those with the lowest velocity thresholds (2°/s) had branches that projected most anteriorly, while those with progressively higher velocity thresholds (up to 200°/s) projected progressively more posteriorly. Afferents that responded to low velocity ramp movements and spiked tonically projected to more posterior areas of the ganglion than those that responded only to movements.Abbreviations A6SCI sixth abdominal sensory commissure I - CO chordotonal organ - DMT dorsal medial tract - G6 sixth abdominal ganglion - LDT lateral dorsal tract - MDT medial dorsal tract - MVT medial ventral tract - R1–4 nerve roots 1–4 - VLT ventral lateral tract - VMT ventral medial tract     

Cider fermentation with three Williopsis saturnus yeast strains and volatile changes

Williopsis saturnus var. subsufficiens NCYC 2728, W. saturnus var. saturnus NCYC 22 and W. saturnus var. mrakii NCYC 500 were used to carry out cider fermentation to assess their impact on the volatile composition of cider. The changes of yeast cell population, °Brix and pH were similar among the three yeasts. Strain NCYC 500 grew best, with the highest cell population of 1.14 × 108 CFU ml−1, followed by strains NCYC 2728 and NCYC 22 (8 × 107 CFU ml−1 and 3.19 × 107 CFU ml−1 respectively). Esters were the most abundant volatiles produced, followed by alcohols. Among the esters, ethyl acetate, 2-phenylethyl acetate, isoamyl acetate, cis-3-hexenyl acetate and hexyl acetate were the major volatiles. The major alcohols were ethanol, isoamyl alcohol, 2-phenylethyl alcohol and isobutyl alcohol. The three Williopsis yeasts transformed volatile compounds during cider fermentation with significant variations in terms of volatile production and degradation. This study implied that fermentation with Williopsis yeasts could result in cider with a more complex yet fruity aroma.     

Olfactory antennal responses to plant volatiles in apterous virginoparae of the vetch aphidMegoura viciae

Electroantennogram (EAG) responses were recorded from apterous virginoparae of the vetch aphidMegoura viciae Buckton (Homoptera, Aphididae) to more than eighty volatile compounds in order to investigate its sensory ability to perceive plant odours. The response profile ofM. viciae reveals a differential sensitivity for the array of plant volatiles tested. The whole group of general green leaf volatiles is very stimulatory. In addition to (E)-2-hexenal, the following compounds of this group elicit large EAG responses: (E)-2-heptenal, 1-octenol-3, hexyl acetate, (Z)-3-hexenyl acetate, hexanol-1, hexanal, 2-heptanone and 3-octanone. Relatively large EAGs are also produced by 4-methoxybenzaldehyde (p-anisaldehyde), hexanonitrile, heptanonitrile, 1,6-hexanedithiol, butyl isothiocyanate, 4-pentenyl isothiocyanate, (−)-(1S)-β-pinene, (+)-(S)-carvone, (−)-(R)-carvone, α-terpineol, linalool and citronellal. The nitriles are the most effective of all plant volatiles tested. Structure-activity relationships occur in various groups of chemicals and members of the green leaf volatiles, benzaldehydes, isothiocyanates and monoterpenes are ranked accordingly. In the group of green leaf volatiles, aliphatic aldehydes are more stimulating than the corresponding alcohols. EAG responses to series of saturated aliphatic alcohols and aldehydes reveal that C₆ and C₇ compounds are the most stimulatory. Dose-response curves show that the rank order of EAG response amplitudes hardly changes at lower dosages. It is concluded thatM. viciae perceives general plant volatiles as well as more-specific components, such as nitriles and isothiocyanates, associated with the odour blends of non-host plant species.     

Aggregation pheromone of Metamasius spinolae (Coleoptera: Curculionidae): chemical analysis and field test

Male Metamasius spinolae (Gylh.) produce several volatile compounds that are likely constituents of its aggregation pheromone. These compounds were identified by volatile collections and gas chromatography (GC), followed by coupled gas chromatography-mass spectrometry (GC-MS), as 2-methyl-4-heptanone [1], 6-methyl-2hepten-4-one [2], and 2-hydroxy-2-methyl-4-heptanone [3]. Preliminary field experiments using synthetic racemates of these compounds showed that significantly more adult cactus weevils were caught in traps baited with the major single compound three or the 2 + 3 binary combination than in unbaited control traps. However, highest trap efficacy occurred with the 1 + 2 binary combination and a blend of all three synthetic compounds plus prickly pear. Potential uses for the cactus weevil pheromone and possible ways to increase trap captures are discussed.     

Pollinator attraction of the wasp-flower Scrophularia umbrosa (Scrophulariaceae)

Certain species of Scrophularia (Scrophulariaceae), such as S. nodosa and S. umbrosa, are mainly pollinated by social wasps and are consequently described as wasp-flowers. Because plants attract their pollinators with the help of various floral cues, such as floral odour and/or optical cues, we have investigated the role of olfactory and visual floral signals responsible for wasp attraction in S. umbrosa. Using a combination of chemical (GC, GC-MS) and electrophysiological analyses (GC-EAD), we identified ten compounds in the complex floral odour bouquet that are detectable by the wasps' antennae. As in the wasp-flower Epipactis helleborine, we found so-called 'green leaf volatiles' (GLVs) in the floral odour; these GLVs are highly attractive to the wasps. GLVs, mostly six-carbon aldehydes, alcohols and acetates, and other volatile organic compounds (VOCs), are emitted by many plants infested with herbivores, e.g. caterpillars. In contrast to other investigated wasp-flowers, behavioural experiments have demonstrated that, in addition to the floral odour of S. umbrosa, visual cues are involved in pollinator attraction.