Phospholipid composition and pheromonal activity of nuptial secretion of the male German cockroach, Blattella germanica

As a part of the sequential courtship behavior of the German cockroach, Blattella germanica, females feed on the nuptial secretion from the male tergal glands. The pheromonal secretion, consisting mainly of oligosaccharides and phospholipids, strongly elicits a feeding response in virgin females. The phospholipids were composed of phosphatidylethanolamine and phosphatidylcholine. Their fatty acid compositions were determined by chromatographic and enzymatic methods. Although an authentic blend of phospholipids (1,2-dioleoylphosphatidylethanolamine and 1,2-dioleoylphosphatidylcholine) showed a marginal phagostimulant activity, addition of the blend to an authentic blend of oligosaccharides (maltose and maltotriose) strongly enhanced the activity to nearly the level of the crude extract. These results indicate that the nuptial feeding behavior is elicited by a synergistic action between phospholipids and oligosaccharides.     

Pheromonal emission during the mating behavior ofEurycotis floridana (Walker) (Dictyoptera: Blattidae)

The sexual behavior of males and females ofEurycotis floridana was investigated and the various associated behavioral sequences are described. Olfactometer data proved that the male produces a volatile sex pheromone attractive at a distance to conspecific females. The male initiates courtship behavior by exposing the glandular areas on the anterior parts of abdominal tergites 2, 7, and 8. This male calling behavior was observed throughout the day. The males can mate when 8 days old, whereas virgin females are sexually receptive 18 days after becoming adults. Once attracted near the male, the female opens her genital atrium and climbs on the back of the male, where she feeds on the glandular secretions that oozed around a little tuft of setae on the first tergite. These setae are mechanoreceptors and they are stimulated when licked by the females, which informs the male that she is in a proper position for copulation.     

Courtship Solicitation by Females of the Barred Sulphur Butterfly (<Emphasis Type="Italic">Eurema daira</Emphasis>) (Lepidoptera: Pieridae)

Virgin females of the subtropical pierid butterfly Eurema daira were observed to actively solicit male courtship. The resulting interactions involved both aggressive contact with male individuals and elaborate female posturing. Cage-based trials showed that there was a direct relationship between the frequency of courtship solicitation attempts and female age. Virgin females under 3 days old showed little or no interest in initiating courtship. By contrast, 23.2 percent of the virgin females over three days old actively solicited available males with peak solicitation behavior exhibited from five day old individuals.     

Courtship and intersexual signaling in the parasitic waspCotesia rubecula (Hymenoptera: Braconidae)

The courtship behavior of the parasitic wasp Cotesia rubeculawas studied in a flight tunnel using standard quantitative ethological techniques. Emission of a female sex pheromone induces searching and signaling behavior in males. Males combine wing-fanning, which produces low-frequency airborne sound, with pulsing behavior, which transmits a vibrational signal through the substrate to the female and induces her receptivity. Female receptivity is indicated by a stereotyped antennal position, which may provide a visual or tactile signal to courting males. Comparison of successful and unsuccessful courtships indicated that courtship success was dependent primarily on the effective production or reception of the male pulse signal. Overall, the sequence of courtship behavior was similar to that reported for other parasitic wasps.     

Courtship inDrosophila sechellia: Its structure,functional aspects,and relationship to those of other members of theDrosophila melanogaster species subgroup

The courtship behavior of Drosophila sechelliais described. Male wing displays are mainly vibration and scissoring, with low levels of rowing. As courtship proceeds the proportion of courtship spent in male wing vibration and licking increases, whereas female movement decreases. The male courtship song of sechelliacontains pulse song but no sine song. This species also shows a distinctive copulation song associated with mounting and copulation. The main cuticular hydrocarbon in females is 7,11-heptacosadiene. The number of copulations increased when flies were placed in the presence of food. Visual and acoustic stimuli appear to be important for mating. A multidimensional comparison was used to compare members of the melanogaster species subgroup, based upon courtship behavior, song characteristics, and cuticular hydrocarbons. A multidimensional comparison of courtship sequences in sechellia, melanogaster, simulans,and mauritianashowed differences in variability between the two island species as compared to the two cosmopolitan species. The courtship song of D. orenais described: it shows both sine and pulse song; there is also a copulation song in this species.     

Courtship in Drosophila quadrilineata with a unique male behavioral element, abdomen bending

This paper describes courtship of Drosophila quadrilineata of the immigrans group and reports a species-specific element, abdomen bending, that has not been described in Drosophila. The male bends his abdomen right and left as a display to the female. During courtship the male circles around the female, bends his abdomen, and taps her. The male licks the female ovipositor repeatedly, often after the females extrusion. Although the females ovipositor extrusion is thought to indicate rejection of a courting male in many species, the male of D. quadrilineata continues courtship. It is possible that the extrusion is a females mating signal to the courting male. The male of D. quadrilineata mounts the female in a rearward position and the genitalia of both sexes couple after mounting. This mounting position is shared by the species belonging to the immigrans group. We never observed behavior in which it appeared as if the male displayed the black stripes to the female. Males court one another and a courtship chain is frequently formed even in the absence of females. The males abdomen bending and the females extrusion followed by the males licking are discussed.     

Effects of male age on the mating behavior of both sexes in the sorghum plant bug, Stenotus rubrovittatus (Hemiptera: Miridae)

In the sorghum plant bug Stenotus rubrovittatus (Matsumura) (Hemiptera: Miridae), mating behavior consists of male courtship and female mating receptivity. Previous studies have found that female age is correlated with ovary development and that mature females with developed ovaries are more receptive to male courtship. Thus, we examined whether male age affects the mating behavior of both sexes and male accessory gland development. Unmated males 0–9 days after emergence and 3-day-old virgin females receptive to male courtship were studied. Immediately after emergence, only 20 % of the males courted females (n = 25). At 3 days old, 68 % of the males courted females (n = 25), the most active age. In contrast, more than 75 % of the courted females were receptive to the male courtship regardless of male age. These results indicate that only male courtship behavior is affected by male age, although it is not enhanced in proportion to male age. Male accessory glands developed with male age. However, no clear relationship was detected between male courtship behavior and accessory gland development. In S. rubrovittatus, it is difficult to explain male courtship behavior solely from male age and accessory gland development.     

Contact Kairomones Mediating the Foraging Behavior of the Aphid Hyperparasitoid Alloxysta victrix (Westwood) (Hymenoptera: Charipidae)

A variety of host-related substances was assessed in relation to their ability to induce arrestment and antennal examination behavior in adult females of the aphid hyperparasitoid Alloxysta victrix (Westwood). In laboratory bioassays, whole-body homogenates of both unparasitized host aphids [Myzus persicae (Sulzer)] and those parasitized by the primary parasitoid Aphidius colemani Viereck failed to elicit a response. However, significant kairomonal activity was associated with aphid cornicle secretions, aphid honeydew, and solvent extracts of parasitized aphids. Solvent extracts of unparasitized aphids (in either hexane or butan-1-ol) failed to elicit arrestment or antennal examination behavior in hyperparsitoid females. The response to aphid honeydew was significantly lower in experienced females than in naive individuals, suggesting that even relatively strong innate responses can be modified by learning.     

Courtship behavior and discrimination between potential mates by maleDelia antiqua (Diptera: Anthomyiidae)

The repertoire of courtship behaviors of male onion maggots,Delia antiqua (Meigen), in a laboratory bioassay chamber, was analyzed by direct observation and by video recordings, in conjunction with a multichannel event recorder. Seven courtship behaviors were categorized: inspection from the substrate, aerial inspection, contact from the substrate, contact from the air, genital alignment, copulation, and male-male interaction. The frequency distribution of copulation bouts was best described by a Poisson distribution; peak mating activity occurred about 1 h into the bioassay. The duration in copulo, however, was extremely variable. On average, males spent 30 sin copulo (n=183); <30% of bouts were >50 s. The ability of males to discriminate between sexes, sexually immature and mature females, and between females ofD. antiqua and the cabbage maggot,Delia radicum (L.), was most pronounced in the elements of genital alignment and attempted copulation. The courtship and mating behavior inD. antiqua is consistent with a sequence that relies initially primarily on indiscriminate visual recognition of a potential mate, followed by species-and sex-specific semiochemical recognition upon contact.     

Structures related to pheromone storage in alar androconia and the female abdominal scent gland of Heliconius erato phyllis,Heliconius ethilla narcaea,and Heliconius besckei (Lepidoptera: Nymphalidae: Heliconiinae)

We describe the morphology of alar androconia and the female abdominal scent gland of Heliconius erato phyllis, Heliconius ethilla narcaea, and Heliconius besckei. Androconial scales of Heliconius, which are arranged in overlapping wing bands, release pheromones during courtship, probably through vibratory movements of male wings over the female to induce her to mate. An antiaphrodisiac is produced by glands located in the valves of the male and is transferred during copulation to the yellow dorsal abdominal sac present in the virgin female, causing this sac to emit a scent that reduces the attractiveness of the female for courtship with other males. Stereomicroscopy, SEM, and TEM analyses were conducted to describe the morphology of the internal and external scales and the external abdominal scent sac. The findings revealed different sizes of external androconial scales and an internal group of porous structural vesicles that are probably related to the preservation of internal space, reception and storage of secretions, and elimination of volatiles when the male is actively involved in courtship. Translucent projections on the female abdominal scent sac create open reservoirs for the reception, storage, and emission of antiaphrodisiac volatiles along with stink clubs. Male valve denticles vary in form and probably attach securely to the female sac during mating, thus ensuring secretion transfer. These features are discussed in the context of a comparative analysis.     

Onion fly,Delia antiqua,oviposition and mating as influenced by insect age and dosage of male reproductive tract extract (Diptera: Anthomyiidae)

One hundred percent of virgin female onion flies,Delia antiqua, receiving 1/20 of a male equivalent of an aqueous extract of mature male reproductive tract remained unmated in the presence of males and began laying unfertilized eggs at a normally mated rate of about 20 eggs/female/day. The 50% behavioral response (BR₅₀) fell between 1/40 and 1/20 of a male equivalent. Sex peptide responses are not always all-or-none. Some females receiving extract at 1/40 male equivalent oviposited at an intermediate rate. Moreover, at low sex peptide dosages, some females were fully activated ovipositionally but were receiptive to mating. A low level of sex peptide was present in 1-day-old males. Sex peptide titer rose with age until plateauing by 6 days posteclosion. Males began mating at 3 days, when they first had ample mature sperm; 50% of 6-day-old males mated. The mean number of females inseminated per male exposed to an excess of virgin females over 24 h was 4.3±0.6 (±SE). Presence of mature eggs was not always a prerequisite for mating, although probability of insemination was correlated with egg maturation. One-day-old preovipositional females receiving 1/20 of a male equivalent of extract began ovipositing when they had mature eggs at 5–6 days old. Therefore, sex peptide may act early and permanently or have a long half-life and affect behaviors once females reach sexual maturity. Male flies provide females with an excess of sex peptide in many cases.D. antiqua males transferred ca. 5–10 times more sex peptide than necessary to activate females fully. We suggest this excess is related to the speed of female response. It is yet unclear whether sex peptide potency or titer in Diptera has become exaggerated by intra- or intersexual selection.     

Diurnal and seasonal changes in the mate-locating behavior of the satyrine butterfly <Emphasis Type="Italic">Lethe diana</Emphasis>

The time schedule of mate-locating behavior in the male satyrine butterfly Lethe diana was studied. Males displayed territorial behavior only in the afternoon, but it was not known whether they attempted to locate mates in the morning. In order for males of most butterfly species to mate with a virgin female, it would be most effective to initiate mate-locating behavior in the morning, when synchronous eclosions occur. Males of L.diana flew patrols in the morning and displayed territorial behavior in the afternoon. The cost of territorial behavior is probably less than that of patrolling, as territorial flights are shorter than those when patrolling. The temporal switching of mate-locating behavior is theoretically favored, since the reproductive value of females decreases later in the day, when males perform the less costly mate-locating behavior. Thus, the diurnal switching of male mate-locating behavior may be an adaptive strategy in response to the rapid decline in the reproductive value of females over the course of a day.     

The role of silk in courtship and chemical communication of the false widow spider, <Emphasis Type="Italic">Steatoda grossa</Emphasis> (Araneae: Theridiidae)

In spiders, sex pheromones are often associated with silk produced by females, and function in mate attraction, recognition, and evaluation. Silk-bound pheromones typically elicit courtship behaviour in web-building spiders. Here we (1) describe courtship interactions of Steatoda grossa males with virgin or mated females, and (2) show that silk and methanol extracts of silk produced by virgin females trigger courtship behaviour (silk production) by males, whereas silk of mated females does not. Our results indicate that (1) virgin females produce a silk-bound sex pheromone, (2) males discriminate between virgin and mated females based on silk cues, and (3) male silk likely functions in sexual communication.     

The nutritional contribution of males affects the feeding behavior and spatial distribution of females in a bruchid beetle,<Emphasis Type="Italic"> Bruchidius dorsalis</Emphasis>

The relationship between the quality/quantity of male investment and the feeding behavior of females was investigated in a bruchid weevil, Bruchidius dorsalis (Fahraeus), whose males donate nutrition via seminal fluid to females. Experiments on the effect of feeding regimes of both sexes on the mating frequency of females showed that females mated at a higher frequency if given low-quality food or poor male investment. On the other hand, the experiment that examined the effect of male investment quality on female feeding behavior showed that females receiving the high-quality investment exhibited feeding behavior less often. These results suggest that male investment and feeding behavior play the same role for B. dorsalis females. These experiments also showed that there are sex-related asymmetries in mating and feeding behaviors: females mated more often but males fed more often. Moreover, a field census suggested that only males visited non-host flowers to feed on the pollen and nectar during the non-flowering period of the host plants; females always stayed on the host plants irrespective of the flowering phenology. These results suggest that in B. dorsalis courtship role reversal and sex-specific feeding modes are fairly fixed and obligatory, and that male investment, derived from sexual selection, could affect the feeding behavior and spatial distribution of both sexes, which may have far-reaching impact in various ecological contexts.     

Virgin male mating advantage in a primitive acoustic insect (Orthoptera: Haglidae)

Male reproductive behavior in the relict flightless haglid, Cyphoderris strepitans,entails the generation of sound signals and the provision of nuptial gifts to mates. These food gifts take two forms: (1) a gelatinous mass (spermatophylax) augmenting the spermatophore and (2) fleshy metathoracic wings adapted to be eaten. The female consumes a portion of the male's underwings during courtship and copulation and the spermatophylax afterward. The incidence of wing-feeding wounds can be used to monitor the mating success of field-caught males. If, when a male mates, he compromises his ability to provide subsequent nutritive gifts, females would benefit by mating with virgin rather than nonvirgin males. To test this, mating success of virgin and nonvirgin males was compared in a field population. Virgins were found to obtain more matings than explained by their numbers relative to nonvirgins in the population. We conclude that, having mated, a male is at a disadvantage, relative to his virgin competitors, in securing further matings.     

Mating behavior of male deer ticksIxodes dammini (Acari: Ixodidae)

To analyze the sexual behavior of male black-legged deer ticks Ixodes dammini,we collected ticks infesting 202 white-tailed deer. On average, 17.7 males and 8.8 females infested each deer. Field-collected males copulated with a mean of 2.25 females, and virgin males mated with 2.4 females. On experimental hosts, males established sexual contact with feeding females and repelled other males, and about half remained paired after their mate detached. Engorged females continue to be receptive, and males mate more readily with them than with nonfed females. We conclude that male I. damminiare endowed with a repertoire of behaviors which favor an opportunistic mating before seeking a host and a preference for mating with feeding females on the host accompanied by tenacious mate guarding.     

Male courtship in Drosophila: the conditioned response to immature males and its genetic control

Experimentally naive male Drosophila melanogaster respond to sexually immature males with intense courtship. However, this response decreases markedly in a short period of time, and "experienced" males then avoid further courtship with immature males for 4 hr. This subsequent inhibition of the courtship response is specific to immature males; the response to virgin females remains intact. This experience-dependent modification in courtship behavior is designated as "conditioned courtship." Seven mutant strains isolated for their inability to express avoidance conditioning (on criteria independent of courtship) were all found to be mutant with respect to expression of conditioned courtship. The potential application of this phenomenon to mosaic analysis of these mutations is posed. Other results indicate that immature males constitutively release a chemical signal that is sufficient for the expression of conditioned courtship. The interpretation of conditioned courtship as a component of fitness is discussed.     

挥发性信息化合物对玉米螟赤眼蜂寄主选择行为的影响

在实验室条件下,利用“Y”型嗅觉仪测定了源于寄主亚洲玉米螟Ostrinia furnacalis (Guenée) 鳞片、卵表以及不同生理阶段的雌蛾附腺的不同浓度的提取物对玉米螟赤眼蜂Trichogramma ostriniae Pang et Chen寄主选择行为的影响,并对其引诱作用大小进行了比较。结果表明： 亚洲玉米螟鳞片的正己烷提取物在0.5 mg/mL和 1 mg/mL浓度时对玉米螟赤眼蜂有明显的引诱作用; 寄主卵的正己烷提取物在5块卵/mL和10块卵/mL浓度时对玉米螟赤眼蜂有显著的引诱作用,而在40块卵/mL浓度时对玉米螟赤眼蜂有极显著的驱避作用; 玉米螟赤眼蜂对亚洲玉米螟交配未产卵和产卵后前期的雌蛾附腺提取液有反应,而对处女蛾和产卵后期的附腺提取液没反应。卵表提取物和附腺提取物比鳞片提取液对赤眼蜂引诱作用强,两者对赤眼蜂的引诱作用没有明显差异。     

同种雄性竞争对手的存在对星豹蛛雄蛛求偶和交配行为的影响

越来越多的研究发现,雄性产生精子(精液)也需付出代价。雄性除了依据配偶质量和竞争对手的竞争强度适应性调整生殖投入外,雄性在求偶和交配行为上也相应产生适应性反应,求偶和交配行为具有可塑性。目前雄性求偶和交配行为可塑性研究主要集中于雌性多次交配的类群中,在雌性单次交配的类群中研究甚少。以雌蛛一生只交配一次而雄蛛可多次交配的星豹蛛为研究对象,比较:(1)前一雄性拖丝上信息物质对后续雄蛛求偶和交配行为的影响,(2)雌雄不同性比对雄蛛求偶和交配行为的影响。研究结果表明,星豹蛛前一雄蛛拖丝上的信息物质对后续雄蛛求偶潜伏期、求偶持续时间和交配持续时间都没有显著影响,但前一雄蛛拖丝上的信息物质对后续雄蛛求偶强度有显著抑制作用。同时,性比对星豹蛛雄蛛求偶和交配行为都没有显著影响。可见,星豹蛛雄蛛对同种雄性拖丝上的化学信息可产生求偶行为的适应性调整,而对性比不产生适应性反应。     

Mating behavior ofPhytoecia rufiventris Gautier (Coleoptera: Cerambycidae)

Sexual behavior between males and females, as well as between males, is described and discussed for the cerambycid beetlePhytoecia rufiventris. The beetles' taxis toward plants taller than average height brings the sexes together from a distance. A male may mount another individual (male or female) and attempt copulation without sex discrimination. The male can discern the sex of another individual only when the terminal part of his abdomen touches the ventral surface of the fifth visible sternite of the latter. No evidence of a sex pheromone is found in this species. Within 1.5–5.5 cm the substrateborne vibrations produced by a moving individual may be the important factor which elicits males to approach a moving individual and attempt copulation. If a female is receptive when a male touches her, he can copulate with her without any courtship display. However, if the female runs away and appears unreceptive, the male will perform courtship displays. Copulation is usually terminated by males. Homosexual behavior between males is discussed.