The role of 1-octen-3-ol as a host-odour attractant for the biting midge, Culicoides impunctatus Goetghebuer, and interactions of 1-octen-3-ol with a volatile pheromone produced by parous female midges

Abstract. 1-octen-3-ol, a component of the body odour of ruminants, is demonstrated to be an attractant for host-seeking, parous female Culicoides impunctatus Goetghebuer (Diptera, Ceratopogonidae).This compound was electrophysiologically and behaviourally active and, in the field, released at 0.11 ± 0.01 mg/day, increased female C.impunctatus catches over a seven-day period compared with control catches.When 1-octen-3-ol was combined either with live female midges or an extract of female midges in an electroantennogram assay, additive effects were recorded.Behaviourally, the same combination resulted in both enhanced attraction and repellency, depending on concentration.Attraction was recorded mainly when sub-threshold concentrations of the separate components were combined.The potential of such a combination as an effective field bait is discussed.     

Electrophysiological and behavioural studies of the biting midge, Culicoides impunctatus Goetghebuer (Diptera, Ceratopogonidae): interactions between some plant-derived repellent compounds and a host-odour attractant, 1-octen-3-ol

Abstract. Electroantennogram (EAG) and y-tube bioassays have been used to demonstrate the repellent properties of five plant compounds with host-seeking parous female Culicoides impunctatus Goetghebuer. The compounds were methyl salicylate and allyl-, butyl-, phenyl- and 2-phenylethyl isothiocyanate. EAG thresholds were 1 times 10^-3 to 1 μg. In the bioassays, maximal repellencies occurred with 1 times 10˜² to lOug. When each compound was combined with 1-octen-3-ol, a confirmed host-odour attractant for C.impunctatus females, additive effects were recorded in EAG assays and in bioassays, all of the compounds either reduced or reversed the attractancy of l-octen-3-ol. Of the isothiocyanates, allyl isothiocyanate was the most potent and when combined with 1-octen-3-ol in field trials, the attractant effect of l-octen-3-ol was reduced.     

Culicoides midge trap enhancement with animal odour baits in Scotland

Examples of the commercial trap Mosquito Magnet Pro (MMP emitting attractant 1-octen-3-ol in carbon dioxide 500 mL/min generated from propane fuel), were run 24 h/day on the Isle of Skye, Scotland, during June-August 2001 and evaluated for catching Culicoides biting midges (Diptera: Ceratopogonidae). From 30 days trapping, the catch averaged 2626 +/- 1358 Culicoides females/trap/day (mean +/- SE, range 558 +/- 139 to 6088 +/- 3597, for five sets of six consecutive nights), predominantly the pest Culicoides impunctatus Goetghebuer (68% overall), plus C. vexans (Staeger) > C. delta Edwards > C. pulicaris (L.) > C. lupicaris Downs & Kettle > C. albicans (Winnertz) > other Culicoides spp. Attempts were made to enhance the odour baiting system by adding hexane-extracts (2.1 mg/day) of hair samples from large host animals, resulting in the following effects on Culicoides collections: sheep - 53 %, red deer - 26 %, calf + 20%, pony + 40%, water buffalo + 262%, with greatest increases for C. impunctatus and C. pulicaris. Serial concentrations of these animal extracts (10(-1) - 10(-3) x 2.2 g/mL) were assayed on parous female C. impunctatus response in a Y-tube olfactometer (air-flow 150 mL/min), and by electroantennogram (EAG) on Culicoides nubeculosus Meigen laboratory-reared parous females. Positive behavioural responses to host odours were dose-dependent: the water buffalo extract being most active (threshold 0.22 g/mL), similar to deer, whereas other host extracts were > or = 10-fold less active. Correspondingly, the EAG threshold was lowest for water buffalo, 10-fold greater for deer, calf and pony, but not detected for sheep. If the active component(s) of these host extracts can be identified and synthesized, they might be employed to improve the capture of Culicoides midges for local control by removal trapping.     

Attraction of male winterform pear psylla to female-produced volatiles and to female extracts and evidence of male–male repellency

Pear psylla, Cacopsylla pyricola (Förster) (Homoptera: Psyllidae), is a major pest of commercial pears in North America and Europe. Olfactometer trials have shown that males of both the summer and winter morphotype are attracted to female-infested host material. Additional work with the summer morphotype has shown that males are attracted to females even in the absence of the host plant, which is evidence that female C. pyricola produce a volatile sex attractant. Here, we describe similar results with the winterform, confirming for this morphotype that the female psylla rather than the infested host material is the source of the attractant. Male winterforms displayed attraction to odors from live females in the absence of the host plant, freshly killed females, and female whole body extracts. The female whole body extracts were at least as attractive as a comparable number of live females, suggesting that we were successful at extracting the components of the attractant with this procedure. All previous olfactometer trials with C. pyricola used the insect as the attractant source; the current study is the first to demonstrate that volatile chemicals isolated from the female insect were attractive to male conspecifics. Winterform males were also assayed to odors produced by conspecific males. We found that male psylla avoided volatile odors from live males, freshly killed males, or whole body extracts of males. To our knowledge, these results are the first indication that males of any member of the Psyllidae avoid odors associated with conspecific males.     

Sex attraction in Bactericera cockerelli (Hemiptera: Triozidae)

The potato psyllid, Bactericera (=Paratrioza) cockerelli (?ulc) (Hemiptera: Triozidae), is a major pest of potato. We examined the role of chemical signals in sex attraction, assessing male and female response to male- and female-produced volatile chemicals. In laboratory olfactometer assays, potato psyllid males were attracted to odorants emitted from live females and from solvent extract of females. These results indicate that the female-produced chemicals responsible for attracting males may be isolated by means of insect extractions. Males were also attracted to volatile chemicals from males and extracts of males, providing the first example of male-male attraction in the Psylloidea. Males exposed simultaneously to odorants from conspecific females and males were preferentially attracted to female odorants, suggesting the presence of a female-specific sex attractant for males. Potato psyllid females avoided volatile chemicals emitted by females and extracts of females and by volatile chemicals emitted by males and extracts of males. Possible explanations for avoidance of conspecifics by females are discussed. This study is the first report of male attraction to volatile chemicals emitted by females and female extracts in the Triozidae and more specifically in the potato psyllid.     

Toward development of neem-based repellents against the Scottish Highland biting midge Culicoides impunctatus

Oil of neem, from the tree Azadirachta indica A. Juss (Meliaceae), was evaluated for repellent and antifeedant activity against Culicoides biting midges (Diptera: Ceratopogonidae), by three complementary methods with serial dilutions. Electroantennograms revealed the sensitivity of Culicoides nubeculosus (Meigen) females to neem > or = 0.10%. Culicoides impunctatus Goetghebuer females were repelled by > or = 1% in a Y-tube olfactometer, Using a membrane feeder for wild-caught parous females of C. impunctatus, the proportion blood-feeding was significantly reduced by topical applications of neem oil > or = 0.10% concentrations, with blood-feeding completely prevented by > or =1%. On the basis of these response data, we developed 2% neem-based formulations for personal protection against biting midges.     

Seasonal emergence, host-seeking activity, age composition and reproductive biology of the mosquito Aedes punctor

Abstract. 1. Seasonal patterns of adult emergence, host-seeking and reproductive ageing in a population of the woodland mosquito Aedes punctor (Kirby) are described from field studies in northern Britain in 1984 and 1985.
2. The pattern of emergence showed little variation between years but duration did vary. Males started emerging before females. Sex ratios at emergence showed a marked female bias. A late, second period of emergence apparently represented a second generation in this normally univoltine species.
3. Host-seeking females were first caught at human bait within a few days of the start of emergence and peak numbers at bait occurred 3–4 weeks later. High numbers lasted for c. 2 weeks. The duration of host-seeking activity was related to weather. The second emergence in 1985 resulted in a second peak of activity in early September and probably extended the flight season.
4. As the season progressed the proportion of females that were parous increased. However, this proportion never reached 1.0, indicating variation in individual reproductive success, which differed between years.
5. The reproductive biology of individuals within the population was investigated. No autogeny was detected, suggesting that the population was entirely anautogenous.     

Development of lactic acid-receptor sensitivity and host-seeking behaviour in newly emerged female Aedes aegypti mosquitoes

The temporal patterns for the development of sensitivity in the lactic acid-excited neurones of the antennal grooved-peg sensilla and for the initiation of host-seeking behaviour by newly emerged virgin female Aedes aegypti mosquitoes were determined and compared. A 1:1 correlation between the presence of a high sensitivity to the host attractant, lactic acid, in the lactic acid-excited neurones and the presence of host-seeking behaviour was observed. This finding supports the notion that changes in the activity of the peripheral sensory system are sufficient, although perhaps not the only means, to control host-seeking behaviour of female mosquitoes.     

A releaser pheromone that attracts males in the urine of mature female masu salmon

A releaser pheromone of masu salmon Oncorhynchus masou was investigated using Y-maze behavioural experiments. During the reproductive season, urine of mature females contains a releaser pheromone which acts as a sex attractant for spermiating mature male parr. The releaser pheromone in mature female urine is one or more low molecular weight substances (less than 10 000) soluble in ether under basic conditions. The attractant was not present in either the coelomic fluid of ovulated females nor in neutral or acidic extracts of female urine which contain free steroids and F-type prostaglandins, respectively.     

Facultative blood-feeding in the crabhole mosquito, Deinocerites cancer

1. Most of the female crabhole mosquitoes Deinocerites cancer Theobald entering bait traps had undeveloped ovaries. 2. Nearly half of the D. cancer females that were captured in bait traps, but denied blood, were gravid following a 1 week holding period; they developed relatively small egg clutches autogenously (mean = 7.4 eggs per gravid female). 3. Although both parous and nulliparous D. cancer females were captured exiting from crabholes, only parous females were attracted to and collected in bait traps. 4. Much larger egg clutches (mean = 41.8 eggs per female) were produced anautogenously by mosquitoes that blood-fed on chickens. 5. All D. cancer females from field collected pupae developed eggs autogenously when allowed to mate and sugar feed in the laboratory. These mosquitoes produced an initial egg clutch with a mean size equivalent to that found in blood-fed, parous mosquitoes from bait traps. 6. In the field, mating and sugar feeding rather than blood-feeding seem to stimulate production of the initial egg clutch. 7. D. cancer blood-feeding appears to be delayed until after the first egg clutch has been laid; some parous crabhole mosquitoes are facultative blood feeders and can produce an egg clutch either with or without a blood meal.     

Responses of the biting midge Culicoides impunctatus to acetone, CO2 and 1-octen-3-ol in a wind tunnel

Responses of host-seeking female Culicoides impunctatus (Goetghebuer) (Diptera: Ceratopogonidae) to acetone, carbon dioxide and 1-octen-3-ol were measured in a wind tunnel. Carbon dioxide, presented as a filamentous plume, increased upwind flight in a dose-dependent manner, up to 0.09% concentration. A homogenous CO2 plume elicited similar upwind responses at concentrations up to 0.09%, whereas higher plume concentrations (> 0.1%) induced erratic responses with a suppression of upwind flight. Bovine equivalent concentrations of acetone (1.5 x 10(-6)g/l) and 1-octen-3-ol (1.3 x 10(-8)g/) failed to induce any significant upwind response when tested alone. In the presence of CO2, however, 1-octen-3-ol showed highly significant increases in upwind responses at concentrations of 1.3 x 10(-1) - 10(-8)g/l. Mixtures of CO2+ acetone also enhanced upwind flight at 1.5 x 10(-9)g/l. High tunnel concentrations of both 1-octen-3-ol and acetone inhibited upwind responses. These findings are discussed in relation to host finding by C. impunctatus and known mechanisms by which upwind flight is initiated and arrested at high odour concentrations.     

Field studies on efficacy of host odour baits for the biting midge Culicoides impunctatus in Scotland

The efficacy of some putative attractants for the biting midge Culicoides impunctatus (Goetghebuer) (Diptera: Ceratopogonidae) was assessed using odour-baited 'delta traps' and suction traps. 1-octen-3-ol was confirmed as a potent olfactory attractant for C. impunctatus when released at 0.06mg/h. Acetone (23mg/h) and a mix of six phenolic compounds (phenol, 3-ethylphenol, 4-ethylphenol, 3-methylphenol, 4-methylphenol and 4-propylphenol), at undetermined release rate, also significantly increased delta trap catches compared to unbaited controls. When tested in combination, there was evidence of synergism between CO2 (0.2L/min) and acetone, 1-octen-3-ol or cow urine, trap catches being, respectively, 4.7, 6.2 and 9.3-fold greater than for CO2 alone. Highest catches were obtained with triple bait combinations comprising cow urine + acetone + CO2 or cow urine + 1-octen-3-ol+CO2, which increased trap catches by X 22 and X 24, respectively, compared to CO2 alone. Culicoides impunctatus was found to be extremely sensitive to CO2 and responses, gauged over two field seasons, showed a significant dose-dependent increase in catch across the entire range of release rates (0.2-2.5 L/min). Responses to these release rates, ranging from small to large mammal equivalents, emphasized the important role of CO2 in host location by C. impunctatus. Uses of olfactory attractants for monitoring and control of Culicoides are reviewed on the basis of these results.     

Manipulation of oviposition patterns of the parasitoidCyzenis albicans (Tachinidae) in the field using plant extracts

We examine the oviposition behavior of the parasitic fly Cyzenis albicans(Fall.) to determine if long-distance or contact chemical cues given off by damaged oak and apple foliage influences the attack of their host the winter moth Operophtera brumata(L.). Wind-tunnel experiments indicate the presence of an attractive odor in oak leaves and the absence of an attractant in apple leaves. The application of oak foliage extracts to apple trees increased the number of parasitoid eggs laid and the level of parasitism of winter moths in the field. The impact of altered host-seeking behavior by the parastoid is discussed in the context of the population ecology of its host.     

Nonsurgical embryo transfer in the scimitar-horned oryx (Oryx dammah): Birth of a live offspring

The objective of this project was to determine if modifications of methods of estrous synchronization, superovulation, embryo recovery, and transfer used successfully in other ungulates, both domestic and nondomestic, could be applied to scimitar-horned oryx (Oryx dammah). Donors were two parous females and recipients were one parous and two nulliparous females that were given a total of two cloprostenol injections at an interval of 0 and 13 or 12 days, respectively. Donors were treated with follicle-stimulating hormone (FSH-P, Schering, Kenilworth, NJ) b.i.d. for 4 days and placed with a fertile male. Seven days after the last FSH-P injection, nonsurgical uterine lavages were performed on both donors. One good-quality embryo at the morula stage was recovered and nonsurgically transferred into the right uterine horn of the parous recipient. A healthy female calf born at 247 days post-transfer represents the first known live birth of scimitarhorned oryx following embryo transfer. These results provide additional evidence that estrous synchronization and embryo transfer techniques used in other ungulates can be applied to endangered antelopes such as the scimitar-horned oryx.     

橘小实蝇植物源引诱活性物质的生物测定

植物源引诱物质可显著提高橘小实蝇Bactrocera dorsalis(Hendel)蛋白饵剂的应用效果,本文测试了柑橘、番木瓜、芒果、番石榴、杨桃的叶片浸提物、果实以及已知植物次生物质对橘小实蝇的引诱效果。植物浸提物的生物测定结果表明,杨桃的二氯甲烷浸提物的引诱效果最佳,平均引诱率为20.83%,对于不同的萃取剂而言,二氯甲烷、乙醇浸提物效果明显优于石油醚,三者浸提的平均引诱率分别为15.67%、15.17%和10.50%;对于不同寄主植物而言,柑橘浸提物的效果最佳,3种有机溶剂浸提物的平均引诱率为18.33%,其中石油醚、二氯甲烷和乙醇浸提物的引诱率分别为20.00%、15.83%和19.17%,其他植物浸提物的引诱效果为杨桃>番木瓜>芒果>番石榴,3种有机溶剂浸提物的平均引诱率分别为17.22%、11.67%、11.11%和10.56%。寄主果实的生物测定结果表明,柑橘的平均引诱率为65.83%,其中雌虫和雄虫的引诱率分别为61.67%和70.00%,明显高于杨桃、番石榴和番木瓜,三者的引诱率分别为31.67%、31.67%和21.67%,与柑橘引诱率差异显著(P<0.05)。已知植物次生物质的生物测定结果表明,甲基丁香酚的引诱效果最佳,平均引诱率为45.00%,其中雄虫和雌虫的引诱率分别为86.67%和3.33%;乙酸乙酯和番石榴香精的引诱效果次之,平均引诱率分别为32.50%和28.33%,其中雌虫的引诱率均为36.67%;杨桃香精和柠檬酸的引诱效果较差,平均引诱率分别为25.00%和26.67%。     

Identification of bloodmeals of the Scottish biting midge, Culicoides impunctatus, by indirect enzyme-linked immunosorbent assay (ELISA)

Abstract. An indirect enzyme-linked immunosorbent assay (ELISA) has been developed to identify the bloodmeal sources of Culicoides impunctatus Goetghebuer. Nightly light trapping in Scotland, between April and September 1992, caught a total of 344 blood-engorged female C. impunctatus , all in May, June and July. The insects were stored in 70% ethanol and tested 5–7 months later. Bloodmeal sources were positively identified for 246 (71.5%). Bovine hosts were most common (38.4%), followed by deer (23.0%) and sheep (9.9%). Only one midge had fed on human blood. Relatively freshly fed (<24h), fully engorged females gave the clearest results. The wider applications of the technique to the study of Culicoides host preferences is discussed.     

The effects of allatectomy and juvenile hormone replacement on the development of host-seeking behaviour and lactic acid receptor sensitivity in the mosquito Aedes aegypti

Host-seeking behaviour in newly emerged Aedes aegypti (L.) females is not expressed immediately after adult eclosion but develops gradually over a period of approximately 3-4 days. This development is accompanied by an apparent maturation of the antennal chemosensory afferent neurons involved in the detection of the airborne host attractant lactic acid. Since these events coincide in time with juvenile hormone-dependent previtellogenic ovarian growth and since the expression of other reproduction-associated behaviour has previously been shown to be dependent on juvenile hormone, the effects of juvenile hormone removal and replacement on the development of host-seeking behaviour and the response characteristics of the lactic acid-sensitive receptors were investigated. No effect of juvenile hormone removal by allatectomy or juvenile hormone replacement or augmentation by topical application of the juvenile hormone mimic methoprene was found. It was concluded that this hormone is not involved in the appearance of host-seeking behaviour or the apparent maturation of the lactic acid receptors that occurs during early imaginal life.     

Female life history and mating patterns among the M group chimpanzees of the Mahale national park,Tanzania

The mating behavior of receptive females in the M group chimpanzees (Pan troglodytes schweinfurthii) of the Mahale Mountains, western Tanzania, was studied by intensive sampling over a period of 5 months. Restrictive matings were observed only between parous females mostly in the ovulatory stage and prime adult males, in particular the alpha. Young, nulliparous and/or non-ovulatory females tended to mate with immature or low-ranking adult males. By contrast, older, parous and/or ovulatory females tended to mate with adult and higher-ranking males. These confirmed the results of a previous extensive study of the same group. In addition, gradual shift from opportunistic to restrictive matings after inter-unit-group transfer was revealed for 2 ex-K-group parous females. The findings were discussed in relation to the life history of female chimpanzees. A National Park since 1984.     

Nest-mate recognition in Manuelia postica (Apidae: Xylocopinae): an eusocial trait is present in a solitary bee

In eusocial Hymenoptera, females are more tolerant towards nest-mate than towards non-nest-mate females. In solitary Hymenoptera, females are generally aggressive towards any conspecific female. Field observations of the nest biology of Manuelia postica suggested nest-mate recognition. Experiments were performed involving two live interacting females or one live female interacting with a dead female. Live females from different nests were more intolerant to each other than females from the same nest. Females were more intolerant towards non-nest-mate than towards nest-mate dead females. When dead females were washed with pentane, no differences in tolerant and intolerant behaviours were detected between non-nest-mate and nest-mate females. Females were more intolerant towards nest-mate female carcasses coated with the cuticular extract from a non-nest-mate than towards non-nest-mate female carcasses coated with the cuticular extract from a nest-mate. The compositions of the cuticular extracts was more similar between females from the same nest than between females from different nests. The results demonstrate for the first time nest-mate recognition mediated by cuticular chemicals in a largely solitary species of Apidae. The position of Manuelia at the base of the Apidae phylogeny suggests that nest-mate recognition in eusocial species apical to Manuelia represents the retention of a primitive capacity in Apidae.     

Costs of and Investment in Mate-Guarding in Wild Long-Tailed Macaques (Macaca fascicularis): Influences of Female Characteristics and Male–Female Social Bonds

Male primates living in multimale groups tend to direct mate and mate-guarding choices toward females of high reproductive value, i.e., high-ranking, parous females, or females with which they share strong bonds. Little is known, however, about the constraints that may limit male mate-guarding choices (the costs of this behavior) and the influence of the females’ quality on male investment in mate-guarding. We aimed to study the effects of female rank, parity status, and male–female social bond strength on the costs of and investment in mate-guarding by males. We carried out our study during two reproductive seasons on three groups of wild long-tailed macaques in Indonesia. We combined behavioral observations on male locomotion and activity with noninvasive measurements of fecal glucocorticoids (fGC). Males spent less time feeding when mate-guarding nulliparous females than when mate-guarding parous females and tended to have higher fGC levels when mate-guarding low-ranking nulliparous females than when mate-guarding high-ranking nulliparous ones. Evolution should thus favor male choice for high-ranking parous females because such a decision brings benefits at proximate (reduced costs of mate-guarding) and ultimate (higher reproductive value) levels. Further, male investment in mate-guarding was flexible and contingent on female reproductive and social value. Males were more vigilant and more aggressive toward other males when mate-guarding females to which they were strongly bonded and/or high-ranking ones than when mate-guarding other females. Our findings bring a new dimension to the study of mate choice by showing that males not only mate preferentially with high-quality females but may also aim to secure paternity with these females through optimized monopolization.