The different effects on courtship of volatile compounds from mated and virgin Drosophila females

Virgin Drosophila melanogaster females, which are courted vigorously, emit pheromones which stimulate males to court each other (Tompkinset al., 1980). Females which have recently copulated are courted less vigrously, and volatile compounds produced by mated females stimulate less courtship between males. Analysis of these compounds from fertilized females by gas chromatography and behavioral assays indicates that mated females emit less of the sex attractant made by virgins and may also produce material which inhibits courtship. These changes in pheromone production are initiated after the first few minutes of copulation.     

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.     

Evidence for acoustic communication in the parasitoid wasp Glyptapanteles flavicoxis

Females of the parasitoid wasp Glyptapanteles flavicoxis (Marsh) (Hymenoptera: Braconidae) deposit sex pheromone on substrate that elicits attraction and wing fanning in conspecific males. We tested the hypothesis that wing fanning sound induces a behavioral response from females which, in turn, affects the males’ orientation toward them. Females exposed to playback of the males’ wing fanning sound engaged in short flights, with sound characteristics different from those of the males’ wing fanning sound. In two‐choice bioassays, playback of the females’ flight sound attracted significantly more males than a silent control stimulus, and in combination with pheromone‐containing body extract of females it attracted more males than female body extract alone. Our data support the conclusion that the males’ wing fanning induces sound and visual reply signals from females that help males orient toward them.     

Evidence for a mate-attracting chemosignal in the dwarf African clawed frog Hymenochirus

Many male frogs and toads possess sexually dimorphic skin glands (breeding glands). However, in most anuran species, the functional significance of the glands is unknown. Here we show that the breeding glands of male dwarf African clawed frogs (Hymenochirus sp. ) release a mate-attractant chemosignal. The mate-attractant activity was assessed using a two-choice aquatic Y-maze. Female Hymenochirus were allowed to choose between different treatment waters (e.g., plain water and water housing males) in the upstream arms of the maze, and the females' movements were monitored by computer-linked motion sensors. Females showed a positive chemotaxis to water housing males and to water containing homogenized breeding glands. Females showed no reaction to water housing conspecific females or to water housing breeding gland-ablated males. Additional choice tests demonstrated that females were more attracted to water housing males than to water housing females and to water containing homogenized breeding glands than to water housing breeding gland-ablated males. Males in the maze showed no response to water housing either females or other males, indicating that the attractant is specific for females and is therefore neither a species aggregation signal nor a food-related attractant. These results represent the first experimental demonstration of a mate-attractant function for anuran breeding glands. Because many anuran species possess breeding glands, these results suggest that pheromonal communication could be more widespread among frogs and toads than previously believed.     

Male indifference to female traits in an arctiid moth (Utetheisa ornatrix)

Abstract.  1. Female Utetheisa ornatrix (Lepidoptera: Arctiidae) mate selectively with large males able to transmit sizeable quantities of nutrient and defensive pyrrolizidine alkaloid with the spermatophore. The female gauges male size indirectly by assessment of the male's courtship pheromone.
2. Male Utetheisa invest upward of 10% of body mass in spermatophore production, and could therefore have been expected to be choosy; however, when offered females differing in alkaloid content, body mass, or mating status, males showed disregard of these parameters, both in their choice of partner and in their allocation of resources to the spermatophore.
3. It is concluded that Utetheisa males do not have the option to select females by comparison shopping . Females broadcast their attractant pheromone for less than an hour per day. Given this time constraint and the potentially high cost of mate localisation, males may have no choice but to mate on an opportunistic basis.     

Foraging behavior of a Dipteran leaf miner on exploited and unexploited hosts

Summary The foraging behavior of females of the leaf miner, Agromyza frontella (Rondani), (Diptera: Agromyzidae) when encountering unexploited or exploited alfalfa plants was studied in large field cages and in laboratory bioassays. Females did not recognize any exploited leaflets before contacting them and did not distinguish between leaflets with an egg or first instar larva and unexploited leaflets, even after contact. Only one fly oviposited in leaflets which contained 80–120 nutrition holes, one late second or third instar larva or which were marked with an epideictic pheromone in field cages. In laboratory bioassays females oviposited less in leaflets containing a second or third instar larva or an empty larval mine than in unexploited ones. Females foraging on unexploited leaflets engaged in area-restricted search and 10 of 11 females remained on the test plant for the full 60 min of observation. However, females foraging on exploited plants were much more active, spent a greater proportion of their time searching for suitable hosts, had the highest rates of visitation to all above ground plant parts and emigrated to the cage walls before 60 min had elapsed. These quantitative measures of foraging behavior indicated that females ranked plants after landing on them in the following order: unexploited plants >plants marked with pheromone or with many nutrition holes >plants with late instar larvae. The order of host ranking by foragers was in general agreement with the suitability of the host plants for larval survival, development and reproduction, as estimated from previous laboratory studies.Females of A. frontella foraging on unexploited alfalfa plants fed and oviposited significantly more often in the upper apical leaflets than in the lower, older leaflets. However, the choice of feeding site by flies on exploited plants did not vary with leaflet position (age), indicating that females fed in order to sample leaflet quality and that females investigated lower (older) leaves after they discovered that the preferred upper leaves were occupied. These data suggest that high quality oviposition sites may be limiting for A. frontella females, which could explain why superparasitism of leaflets sometimes occurs in nature, even when unexploited sites are available.     

Bioassay evidence for a sex attractant pheromone in the large carpenter bee, Xylocopa varipuncta (Anthophoridae: Hymenoptera)

A field bioassay was conducted to test i˜hethercompoundsp roduced in the thoracic gland of male X. varipuncta are involved in long-range mate attraction. Filter paper impregnated with the two most abundant compounds of the male gland was placed in sites comparable to those used by territorial males during the time of day when mating activity occurs. Females visited these filter paper 'lures' as often as they visited unmanipulated males in the study area. Males with the chemical mixture added to their territories did not receive more matings than males at territories where we had not added the chemical mixture. These findings and other studies, summarized herein, support the hypothesis that the male gland produces a pheromone which functions as a long-range sex attractant.     

Grasshopper sexual pheromone: a component of the defensive secretion in Taeniopoda eques

ABSTRACT. Both male and female Taeniopoda eques (Burmeister) emit a defensive secretion from their thoracic spiracles. The secretion from mature females was found to act as a sex pheromone, eliciting mating behaviour in males. Females became chemically attractive to males about 16–18 days after eclosion. The antennae are shown to be the site of pheromone reception in males. Feeding upon natural host plants was not a prerequisite for pheromone production.     

Modeling intra-sexual competition in a sex pheromone system: how much can female movement affect female mating success?

Mating disruption theory predicts that high concentrations of female pheromone, and/or large numbers of release sites, should confuse males orienting to "calling" females, reduce the number of successful matings, and decrease the reproductive potential of the population. In this scenario, females are regarded as stationary point sources of pheromone. Past behavioral observations, however, have shown virgin female grape root borers, Vitacea polistiformis Harris, significantly alter their behavior in mating disruption treatments. Treated females call at different heights, move less before call initiation, and move more after call initiation than control females. Pheromone gland dragging and wing fanning also increase significantly during pheromone treatments. These behavioral differences are significant only if they alter the mating success of females. Because long-term field studies are impractical, we used known behavior of male and female GRB to build a Fortran language time step model, adding the effects of female movement to past models of male pheromone plume following. Females were distributed randomly, and then assigned a conditional movement strategy. If females were within the competitive portion of another female's plume, the downwind female moved. Except in the lowest population density tested, females moving upwind and crosswind when in a competing female's pheromone plume mated significantly more often than females remaining stationary. In all population simulations, mating success was significantly reduced when females moved downwind. These field and simulation studies provide strong evidence for female movement as a previously overlooked potential mechanism for resistance to mating disruption treatments, as well as a shaping behavior in the evolution of pheromone communication systems.     

Effects of decapitation and PBAN injection on amounts of triacylglycerols in the sex pheromone gland of Manduca sexta (L.)

The correlation between triacylglycerols containing conjugated diene fatty acyl moieties and pheromone aldehydes in the sex pheromone glands of females of Manduca sexta was investigated. Females decapitated 15 h after adult emergence neither called nor produced pheromone during the natural period of pheromone production on the subsequent two nights. However, these females could be stimulated to produce sex pheromone for prolonged periods by repeated injection of synthetic pheromone biosynthesis activating neuropeptide (PBAN). Gas chromatographic analysis of methanolysis products of lipids extracted from the pheromone glands of decapitated and intact females showed no differences in the amounts of fatty acyl precursors of pheromone. High performance liquid chromatographic analysis of the triacylglycerols containing conjugated diene analogues of the pheromone components (diene TG), obtained 24 and 48 h after decapitation, showed that the total amounts of these components were not affected by decapitation. The amounts of all diene TG peaks declined significantly when decapitated females were stimulated to produce pheromone during a 7 h period by repeated injection of PBAN at 3 h intervals but recovered when pheromone production subsided. These results indicate that PBAN induces liberation of pheromone precursors from the triacylglycerols during pheromone biosynthesis but does not induce replenishment of this storage pool. © 1996 Wiley-Liss, Inc.

1 This article is a US Government work and, as such, is in the public domain in the United States of America

     

Determination of the amount of sex pheromone emitted by individual virgin females of the white grub beetle, <Emphasis Type="Italic">Dasylepida ishigakiensis</Emphasis> (Coleoptera:Scarabaeidae) at different stages of reproductive life

Virgin females of the white grub beetle, Dasylepida ishigakiensis Niijima et Kinoshita, emit the sex pheromone, 2-butanol. To investigate the relationships between pheromone emission and ovarian development in this beetle, virgin adult females were kept at 18°C for various lengths of time. The sex pheromone released by individual females at first calling was collected at 23°C and the amounts determined. Females started developing visible oocytes at day 44 and emitting sex pheromone at day 50. Almost all females at day 52 or later possessed eggs, indicating that diapause had ended. A high positive correlation was found between the amounts of pheromone emitted and levels of ovarian development. A large variation of pheromone emission was encountered even among individuals of the same age. Females retained the ability to emit sex pheromone as long as they were virgin. These results may provide important information regarding the timing and duration of application of synthetic sex pheromone in an attempt to control this pest through disrupting the male-female communication.     

Predation risk and the opportunity for female mate choice in a coral reef fish

The influence of predation risk on the opportunity for female mate choice was investigated in the tailspot wrasse Halichoeres melanurus at two sites on a coral reef in Okinawa, Japan. Females mated repeatedly with the nearest males, but they also changed mates frequently at both sites. Mate changes were seen not only in the context of spatiosocial changes (mate disappearance or shifts in male territories) but were also probably the result of actual mate choice by females. Females at one site (site A) changed mates more often and conducted longer spawning trips from their home ranges to male territories than at the other site (site B). Fish at site A were faced with a higher frequency of predators than that of site B. However, fish of site A suffered fewer attacks from predators because they had more shelter, suggesting lower predation risk in that site. These results suggest that females under higher predation risk had less opportunity to choose preferable mates and that they had to mate with the same, nearest males in most cases.     

A density dependent model of pheromone trapping

A discrete, deterministic model is presented to represent a use of pheromones in insect pest control. Males are assumed to be attracted to traps containing virgin females releasing male-attracting pheromone (or equivalent pheromone preparation); males entering the traps are killed. Females are assumed monogamous, and males can mate only once per day; both sexes have the same density dependent mortality. If the amount of pheromone is above a certain value, so that males become limiting, there are two positive equilibria; the larger is locally asymptotically stable, thus control becomes possible. As the amount of pheromone is further increased, these two equilibria merge at a critical release rate, above this value the pest is eradicated. Density dependence makes eradication by this means much easier, for parameter values considered this critical release rate is about two orders of magnitude less than that in a density independent case.     

Laboratory response of female Lutzomyia longipalpis sandflies to a host and male pheromone source over distance

A nylon net cage (260 x 40 x 40 cm) provided with unidirectional air flow was used to demonstrate the response of female sandflies, Lutzomyia longipalpis Lutz & Neiva (Diptera: Psychodidae) to a host and extract of male pheromone glands over a distance of 240 cm. Females responded more rapidly and in greater numbers to the host if male pheromone was present. Results suggest that one function of the pheromone may be to attract females from some distance to a common mating site, on or near the host.     

(1S,3S,7R)-3-methyl-alpha-himachalene from the male sandfly Lutzomyia longipalpis (Diptera: Psychodidae) induces neurophysiological responses and attracts both males and females

Lutzomyia longipalpis adult males form leks on or near hosts and release (1S,3S,7R)-3-methyl-alpha-himachalene from their tergal glands to lure females to the same site for mating and feeding. Here we have examined whether the male-produced attractant could also serve as a male aggregation stimulus. High resolution chiral capillary gas chromatography analysis of male tergal gland extracts, synthetic (1S,3S,7R)-3-methyl-alpha-himachalene, and a synthetic mixture of all isomers of 3-methyl-alpha-himachalene, was coupled to electrophysiological recordings from ascoid sensillum receptor cells in antennae of male and female sandflies. Receptor cells of both sexes responded only to the main component of the male tergal gland extract that eluted at the same retention time as (1S,3S,7R)-3-methyl-alpha-himachalene. Furthermore, of the eight 3-methyl-alpha-himachalene isomers in the synthetic mixture only the fraction containing (1S,3S,7R)-3-methyl-alpha-himachalene, co-eluting with an isomer of (1S*,3S*,7S*)-3-methyl-alpha-himachalene, elicited an electrophysiological response from male and female ascoid sensillum receptor cells. Both males and females flew upwind in a wind tunnel towards a filter paper disk treated with either 4-6 male equivalents of the tergal gland extract, pure (1S,3S,7R)-3-methyl-alpha-himachalene or the synthetic mixture of eight isomers. This indicates that (1S,3S,7R)-3-methyl-alpha-himachalene derived from L. longipalpis males may have a dual function in causing male aggregation as well as serving as a sex pheromone for females.     

Pheromone pre-exposure and mating modulate codling moth (Lepidoptera: Tortricidae) response to host plant volatiles

Abstract 1 Two codling moth Cydia pomonella kairomonal attractants, ethyl (E,Z)‐2,4‐decadienoate (pear ester) and (E)‐β‐farnesene, were tested in an insecticide‐sprayed apple orchard and an orchard treated for mating disruption with synthetic pheromone (E,E)‐8,10‐dodecadienol (codlemone). Male captures with pear ester were higher in the pheromone‐treated than in the insecticide‐treated orchard, whereas captures with (E)‐β‐farnesene were not different. Subsequent wind tunnel experiments confirmed that pre‐exposure to sex pheromone codlemone increased the behavioural response of codling moth males to pear ester. This supports the idea that male attraction to the plant volatile pear ester and sex pheromone codlemone is mediated through the same sensory channels. 2 Pear ester is a bisexual codling moth attractant and even captures of female moths were significantly increased in the pheromone‐treated orchard. In the laboratory wind tunnel, pheromone pre‐exposure had no effect on female response to pear ester, but significantly more mated than unmated codling moth females flew upwind towards a pear ester source. Differences in mating status in insecticide‐treated vs. pheromone‐treated orchards may thus account for the differences in female trap captures with pear ester. 3 These findings are important with respect to monitoring of codling moth with pear ester in mating disruption orchards. They also emphasize the importance of host plant volatiles in pheromone‐mediated mating disruption, which has been neglected to date.     

Semiochemical emission from individual galleries of the southern pine beetle, (Coleoptera: Curculionidae: Scolytinae), attacking standing trees

We collected, identified, and quantified volatiles arising from individual gallery entrances of the monogamous bark beetle Dendroctonus frontalis Zimmermann. Samples were collected while the insects were mass attacking mature loblolly pines (Pinus taeda L.) in an established infestation in western Mississippi, 1 August through 3 October 2005. Following volatile sample collection, the entrances were dissected and categorized according to those that 1) contained a solitary female (the gallery initiating sex), 2) contained a pair that had not yet produced an egg gallery, 3) led to an egg gallery with niches and/or eggs, or 4) represented failed attacks (either abandoned or containing dead beetles). The greatest mean release rate of the female-produced aggregation pheromone components frontalin (74 ng/h) and trans-verbenol (0.35 microg/h) was detected from entrances of solitary females, whereas the highest mean quantities of the male-produced multifunctional pheromone components endo-brevicomin (18 ng/h) and verbenone (0.15 microg/h) were detected from entrances of preoviposition beetle pairs. Alpha-pinene, a host-produced monoterpene that functions as a synergist for the aggregation attractant for D. frontalis, was detected from entrances of solitary females and preoviposition pairs at a rate of 0.6 mg/h, or 3-4 orders of magnitude greater than the insect-produced components of the attractant. Our results indicate that the release rates of pheromone components used in published field studies of the chemical ecology of D. frontalis (generally > 0.1 mg/h) represent thousands of 'attack equivalents' or production rates on the scale of a beetle mass attack on a single host. Additionally, our data suggest that the loss in attractiveness of host tissue fully colonized by D. frontalis is because of the disappearance of attractants rather than an increase in inhibitors.     

Effects of exposure to pheromone and insecticide constituents of an attracticide formulation on reproductive behavior of oriental fruit moth (Lepidoptera: Tortricidae)

The effect of exposure to both the pheromone and insecticide constituents of an attracticide formulation on subsequent pheromonal response of male oriental fruit moth, Grapholita molesta (Busck) (Lepidoptera: Tortricidae), was tested in several wind tunnel bioassays. Male response to the attracticide formulation was significantly reduced in all behavioral categories, including source contact 1 h after sublethal exposure (both by voluntary contact in the wind tunnel and forced application in the laboratory) to the attracticide formulation containing inert ingredients, pheromone, and insecticide. Sublethal exposure to the attracticide formulation in the laboratory (forced application) 24 h before the bioassay resulted in a significantly lower proportion of males subsequently responding to attracticide droplets in the wind tunnel. However, voluntary contact of male moths with the toxic formulation in the wind tunnel had no effect on subsequent response 24 h later. Exposure of males to different constituents of the attracticide formulation demonstrated that both pheromone and insecticide exerted effects on subsequent male pheromonal response. Exposure to the formulation containing the inert ingredients plus the pheromone (no insecticide) significantly reduced male behavioral responses to an attracticide droplet in the wind tunnel 1 h but not 24 h after exposure, compared with males treated with inert ingredients alone. Response to attracticide droplets was further reduced by exposure to the entire attracticide formulation containing inert ingredients, pheromone and insecticide at both 1 and 24 h postexposure. Similarly, males exposed to inert ingredients plus pheromone were less likely to orient to female-produced plumes 1 h but not 24 h after exposure than males treated with inert ingredients alone. Response to female-produced plumes was further reduced at 1 h but not at 24 h after exposure to the entire attracticide formulation. Mating success of males was significantly reduced by exposure to the entire attracticide formulation but not to the formulation without insecticide when placed with females 1 and 24 h postexposure. These findings suggest that sublethal poisoning of males exposed to the attracticide formulation will enhance the effectiveness of this formulation under field conditions.     

The responses ofPraon spp. parasitoids to aphid sex pheromone components in the field

In autumn 1991, aphid parasitoids of the genusPraon (Hymenoptera; Braconidae) were caught in water traps with lures containing synthetic aphid sex pheromone components at three sites in England and one in Germany. At two of the English sites and at the German site, the traps were placed in winter cereal fields whilst the third English site was in woodland. Three species were caught,P. volucre, P. dorsale andP. abjectum. Those caught in cereal fields were almost entirelyP. volucre, whilstP. dorsale dominated at the woodland site. Of the known aphid sex pheromone components, the most effective lure was the (+)-(4aS, 7S, 7aR)-nepetalactone. Nepetalactone traps placed at the woodland site in spring and summer caught fewPraon females, and attraction may be confined to the autumn, when sexual female aphids are present in the field. Male parasitoids did not respond to the aphid pheromones at any time, although they were caught in suction traps operated at the woodland site during the autumn. At the cereal sites height had a significant influence on the efficiency of the pheromone traps, those placed just above the crop canopy being most effective. There was no evidence that any other genus of parasitoid responded to aphid sex pheromones at these sites.     

Effects of age, diet, female density, and the host resource on egg load in Anastrepha ludens and Anastrepha obliqua (Diptera: Tephritidae)

Oocyte counts, used as a measure of egg load, were compared among three different age groups (15, 30 and 45 days) of two polyphagous species of tephritid fruit flies, Anastrepha ludens and Anastrepha obliqua, which were exposed to varying conditions of diet (sucrose vs sucrose and protein), availability of oviposition substrate (present vs absent), adult female density (1, 2 and 4 females/cage), and semiochemical context (presence vs absence of male pheromones and fruit volatiles). In both species, oocyte counts were higher in older females and for females fed sucrose and protein than for females fed sucrose only. The presence of artificial oviposition substrates influenced oocyte counts in A. obliqua, but not in A. ludens. Female density influenced oocyte counts in both species. Females maintained in groups had higher egg loads than isolated females. Finally, preliminary evidence suggests that semiochemical context influenced oocyte counts. Counts were highest for females in a room containing both fruit volatiles and male pheromone, lowest for females in a room containing neither volatiles nor pheromone, and intermediate for females in rooms containing either volatiles or pheromone but not both. Our results suggest that egg load is influenced by environmental factors in different ways in these two species. Egg load in A. obliqua, a species whose host fruits are highly ephemeral, is responsive to access to the host resource. By contrast, in A. ludens, a species infesting less ephemeral fruit, female density and age played a more important role than host stimuli. The role of ovarian maturation and oviposition in mediating these effects, as well as implications for mass rearing and pest management, are discussed.