Genetic and endocrine studies of the pregnancy-blocking pheromone of mice.

Secretion of the pregnancy-blocking pheromone was stimulated by injection of depo-testosterone cypionate into females and males of inbred strains of mice which do not normally secrete the pheromone. Testosterone treatment of SJL males altered pheromone secretion so that pregnancies were blocked when the stud male was of the same inbred strain; an event that does not normally occur. Injection of epiandrosterone, androstenedione, androsterone or testosterone significantly increased pheromone secretion in SJL females, but progesterone and dehydroepiandrosterone were ineffective. Kidney weights were significantly increased by administration of androgen metabolites and the possibility of the kidney being the site of pheromone synthesis is discussed. The preputial gland can be excluded as the site of pheromone synthesis since males which are hemizygous for the Tabby-J gene and have no preputial glands blocked pregnancies as effectively as their normal littermates. Preliminary results are also presented concerning the isolation of the pregnancy-blocking pheromone from urine. Urine was analysed by gas chromatography and a peak was observed whose concentration could be correlated with secretion of the pheromone, although the compound(s) has not been identified or tested for biological activity.     

Age-dependent changes in structure and function of the male labial gland in Bombus terrestris

The cephalic region of the labial gland in the buff-tailed bumblebee, Bombus terrestris, consists of numerous acini (formed by associated secretory cells and a central lumen) and connecting ducts. Age-dependent changes in secretion production (both qualitative and quantitative) are associated with changes in the amount of rough endoplasmic reticulum (RER), Golgi apparatus, and smooth endoplasmic reticulum (SER). The main secretory organelle is RER in the youngest individuals (pharate, and less-than-a-day old males), Golgi apparatus in 1-day-old males, and SER in males older than 2 days. Secretory cell death starts at 5 days of age, with maximal longevity to 10 days. Pheromone production starts immediately after eclosion, with pheromone quantities increasing until day 7. 2,3-dihydrofarnesol, the main component of the male-marking pheromone, appears in 1-day-old male glands, and reaches a maximum at 7 days of age, when its presence in the gland starts to decrease gradually. Older glands contain compounds not present in young ones. Variation in pheromone quantity and composition are reflected sensitively in the response of the queen antennae. Though queen antennae responded to gland extracts of all ages examined, maximum sensitivity was observed in response to extracts of glands 2-10 days old, while extracts of older glands gradually lose their effectiveness. Both major and minor components of the labial gland secretion extract elicited queen antennal responses suggesting that the pheromone is a multicomponent blend. Age-dependent changes in pheromone production, accumulation and tuning of pheromone activity are all synchronized approximately with male flight from the hive.     

Spermatophore formation in the pseudoscorpion Chthonius ischnocheles (Chthoniidae)

The reproduction of pseudoscorpions involves indirect sperm transfer by means of spermatophores. The spermatophores are the product of the male genital atrium. A functional interpretation of spermatophore formation in Chthonius ischnocheles is based on evidence from (a) a morphological study of the genital atrium, the associated accessory glands and the musculature (b) males sectioned during spermatophore production (c) histochemical tests on the glands and their secretions (d) biochemical analyses of one gland (posterior dorsal) and its secretion (e) the behaviour of males during this process and (f) the structure of the spermatophore.
The anterior region of the genital atrium is concerned with the production of the sperm packet. The encysted sperm and the seminal fluid from the prostatic reservoir are encapsulated in a sperm packet by a secretion from two pairs of anterior glands. The posterior region of the genital atrium is responsible for the formation of the spermatophore stalk and its distal elaboration, the two lateral collars. These parts of the spermatophore arise from the fibroin secretion of the posterior dorsal gland; the shape of the spermatophore collars is correlated with their mould, the medial diverticula. In addition the lateral glands secrete a light oil which accumulates on a thickening of the spermatophore stalk proximal to the collars. This suggested that this secretion acts as a pheromone to attract females to the spermatophore since in this species males produce their spermatophores in the absence of females.     

11-ketotestosterone stimulates putative sex pheromone production in the male peacock blenny, Salaria pavo (Risso 1810)

Male peacock blennies (Salaria pavo) release odors from their anal glands and, possibly, from their gonads that attract reproductive females. The current study investigated the effects of 11-ketotestosterone (KT) on development of the anal glands, testes, and accessory testicular organs as well as the subsequent olfactory potency of their secretions. After 3 wk of KT treatment (5 mg/kg in silastic implants), clusters of cells secreting neutral mucins differentiated in the anal gland of all treated males, whereas this occurred in only one control male. Secretions by anal glands from KT-treated males elicited greater olfactory responses, as assessed by electro-olfactography, than those from controls. Treatment with KT stimulated testicular gland growth and sialomucin secretion but had no clear effect on the germinal region of the testis; KT also stimulated enlargement of, and fluid secretion in, the blind pouches (paired evaginations of the spermatic ducts). Secretions by the testes and fluid in blind pouches from KT-treated males elicited greater olfactory responses than those from controls. In conclusion, KT stimulates development of the anal glands and testicular accessory organs and promotes odorant secretion, the putative multicomponent male peacock blenny pheromone.     

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.     

Cuelure but not zingerone make the sex pheromone of male Bactrocera tryoni (Tephritidae: Diptera) more attractive to females

In tephritid fruit flies of the genus Bactrocera Macquart, a group of plant derived compounds (sensu amplo ‘male lures’) enhance the mating success of males that have consumed them. For flies responding to the male lure methyl eugenol, this is due to the accumulation of chemicals derived from the male lure in the male rectal gland (site of pheromone synthesis) and the subsequent release of an attractive pheromone. Cuelure, raspberry ketone and zingerone are a second, related group of male lures to which many Bactrocera species respond. Raspberry ketone and cuelure are both known to accumulate in the rectal gland of males as raspberry ketone, but it is not known if the emitted male pheromone is subsequently altered in complexity or is more attractive to females. Using Bactrocera tryoni as our test insect, and cuelure and zingerone as our test chemicals, we assess: (i) lure accumulation in the rectal gland; (ii) if the lures are released exclusively in association with the male pheromone; and (iii) if the pheromone of lure-fed males is more attractive to females than the pheromone of lure-unfed males. As previously documented, we found cuelure was stored in its hydroxyl form of raspberry ketone, while zingerone was stored largely in an unaltered state. Small but consistent amounts of raspberry ketone and β-(4-hydroxy-3-methoxyphenyl)-propionic acid were also detected in zingerone-fed flies. Males released the ingested lures or their analogues, along with endogenous pheromone chemicals, only during the dusk courtship period. More females responded to squashed rectal glands extracted from flies fed on cuelure than to glands from control flies, while more females responded to the pheromone of calling cuelure-fed males than to control males. The response to zingerone treatments in both cases was not different from the control. The results show that male B. tryoni release ingested lures as part of their pheromone blend and, at least for cuelure, this attracts more females.     

Response of workers of Atta sexdens rubropilosa (Hymenoptera: Formicidae) to mandibular gland compounds of virgin males and females

Abstract.  The secretion from the mandibular glands of males of the leaf-cutting ant Atta sexdens rubropilosa is responsible for the reaction of workers outside the nest at the time of sexual swarming. Workers respond with excitability and aggression when presented with the natural mixture of 4-methyl-3-heptanol and 4-methyl-3-heptanone, which is contained in the secretion of the male mandibular glands. Workers respond quickly to fractional amounts of one male equivalent. 4-Methyl-3-heptanone, from the virgin female mandibular glands causes much less response in workers, whereas an equimolar mixture of male and female pheromones gives a still less clear response. The male pheromone plays the most important part in the communication of workers outside the nest at this time.     

Male moth songs tempt females to accept mating: the role of acoustic and pheromonal communication in the reproductive behaviour of Aphomia sociella

Background

Members of the subfamily Galleriinae have adapted to different selective environmental pressures by devising a unique mating process. Galleriinae males initiate mating by attracting females with either chemical or acoustic signals (or a combination of both modalities). Six compounds considered candidates for the sex pheromone have recently been identified in the wing gland extracts of Aphomia sociella males. Prior to the present study, acoustic communication had not been investigated. Signals mediating female attraction were likewise unknown.

Methodology/Principal Findings

Observations of A. sociella mating behaviour and recordings of male acoustic signals confirmed that males initiate the mating process. During calling behaviour (stationary wing fanning and pheromone release), males disperse pheromone from their wing glands. When a female approaches, males cease calling and begin to produce ultrasonic songs as part of the courtship behaviour. Replaying of recorded courting songs to virgin females and a comparison of the mating efficiency of intact males with males lacking tegullae proved that male ultrasonic signals stimulate females to accept mating. Greenhouse experiments with isolated pheromone glands confirmed that the male sex pheromone mediates long-range female attraction.

Conclusion/Significance

Female attraction in A. sociella is chemically mediated, but ultrasonic communication is also employed during courtship. Male ultrasonic songs stimulate female sexual display and significantly affect mating efficiency. Considerable inter-individual differences in song structure exist. These could play a role in female mate selection provided that the female''s ear is able to discern them. The A. sociella mating strategy described above is unique within the subfamily Galleriinae.     

The ultrastructure of a pheromone-secreting gland in the male scorpion-fly Harpobittacus Australis (Bittacidae: Mecoptera)

The eversible vesicles of the sex pheromone glands consist of three cell types, secreting the cuticle, the duct and the pheromone, respectively. Each pheromone-secreting cell has an end apparatus, bounded by microvilli and lined by a granular mass, that is penetrated by remarkable elongate filaments extending far up into the secretory duct. Vesicles containing secretion or secretion precursors abound. The possible role is discussed of the various organelles in the elaboration and transport of the pheromone.     

The morphology of the sex pheromone gland in Drosophila grimshawi

The morphology of a sex pheromone-producing gland found in the abdomen of Drosophila grimshawi males was studied by light and electron microscopy. This gland, consisting of two intra-anal lobes, contains cells that resemble those of other insect pheromone glands. However, in contrast to many other insect pheromone glands that release pheromone through the cuticle, cells of the intra-anal lobes secrete into a canaliculi-duct system that empties into the anal region. The liquid secretory product flows along the surface of the intra-anal lobes and is brushed onto the substrate by fingerlike projections on the lobes' surfaces.     

榆木蠹蛾性信息素通讯系统的超微结构观察

为了解榆木蠹蛾Holcocerus vicarius （Walker）雌蛾性信息素分泌腺（性信息素释放系统）位置、 表面形态和超微结构及雄蛾触角感受器（性信息素接收系统）的种类、 形态、 分布及功能, 利用扫描电镜和透射电镜对榆木蠹蛾雌蛾性信息素分泌腺和雄蛾的触角进行观察。结果表明： 榆木蠹蛾雌蛾性信息素分泌腺位于腹尖末端第8～9节节间膜上的背面中央区域, 腺体表面分布着许多饱满的锥状突起, 2日龄处女雌蛾腺体细胞间有明显的胞连接, 细胞基底膜基褶较多, 质膜上分布着微绒毛, 并与内表皮连接, 内表皮上含有多层几丁质, 胞质中含有脂质粒、 大量空泡、 光面内质网、 粗面内质网及线粒体; 雄蛾触角鞭节上有5种感受器, 为毛形感器、 刺形感器、 锥形感器、 腔锥形感器和曲毛形感器, 其中毛形感器数量最多, 曲毛形感器最少。柄节和梗节被大量鳞片覆盖, 未观察到感器。榆木蠹蛾性信息素通讯系统的研究为榆木蠹蛾性信息素的生物合成、 性信息素的提取、 鉴定及成虫生殖交配生物学行为提供了依据。     

The scent glands of the male south american locust Schistocerca cancellata, an electron microscope study

The male South American locust, Schistocerca cancellata, emits a strong aromatic scent at the time of maturation. This aroma is characteristic of mature adult males living in crowded conditions. In isolated males and in females it is perceptible but faint, while nymphs emit no scent at all. Since dermal glands are numerous in the mature adult but much rarer in nymphs, young imagos and females, it seems likely that the scent is associated with these glands. The scent seems to be a maturation-pheromone, stored and released by these dermal glands. Each gland consists of a gland cell and a duct cell (type 3 gland in the classification of Noirot and Quennedey, (1974)). Each gland cell has a single end-apparatus consisting of an oblong cavity limited by the projecting tips of densely packed microvilli. A duct, dilated to form three successive bulbosities at its distal end, opens in the cavity from which it conveys the secretion to the outside. A network of fibrillar material anchors the duct to the cavity. Reproductive synchrony, beneficial in social insects, seems to be achieved in locusts by the aromatic pheromone released by the glands at the time of maturation.     

Two body fluids containing bovine estrous pheromone(s)

Bovine females produce an estrous pheromone(s). Because onlyother cattle are reliable detectors of the pheromone(s), weevaluated various body fluids (cervico-vaginal mucus, serumand vulval skin gland secretion) with a bull behavioral assayin the absence of a female. It specified 15 behaviors involvingattraction to the body fluids, detection of the stimulus, andsubseqeunt sexual preparation responses. These behaviors werestereotypical and occurred in a specific sequence. We discoveredthat serum (10–15 ml) taken at estrus was behaviorallypositive, as well as vulval skin glands secretion (<1 mlon a swab, dissolved in 5 ml distilled water). We postulatethat vulval skin glands may be a specialized site for the productionor concentration of the pheromone or that the blood transportsthe pheromone(s) either from vulval skin glands or another source.     

四川短尾鼩包皮腺的挥发性化学成分分析

哺乳动物的包皮腺分泌物对个体间性引诱及繁殖行为的信息交流和种群调节有着重要的作用。本研究以四川短尾鼩（Anourosorex squamipes）为研究对象对劳亚食虫目动物包皮腺挥发性化学成分和化学通讯功能进行报道。采用顶空固相微萃取—气质联用（HS-SPME-GC/MS）的方法,分析四川短尾鼩雄体包皮腺中挥发性物质的化学组成。结果表明：（1）四川短尾鼩雄体包皮腺的挥发性化学成分主要含烷烃类、醇类、酮类、醛类、醚类、酯类、酸类、芳香烃类等45种化合物;（2）成年雄性四川短尾鼩的包皮腺中挥发性成分有39种,幼年雄性含有28种,说明四川短尾鼩成年雄体的包皮腺中挥发性化学成分多于幼年雄体;（3）成年雄体的包皮腺分泌物中含有4种特有的挥发性化学成分,幼年雄体中特有成分仅为1种,表明四川短尾鼩包皮腺中的挥发性化学成分存在年龄差异;（4）四川短尾鼩包皮腺分泌物中不同化学成分相对含量不同,同种化学成分在不同个体间的相对含量存在差异。四川短尾鼩雄性包皮腺挥发性化合物种类丰富,个体、年龄差异明显。推测挥发性化合物中丁酸（Butanoicacid）、乙酸乙酯（Ethyl acetate）、苯酚（Phenol）为四川短尾鼩的信息素。本研究为进一步验证该物种的信息素成分及其传递机制提供了基础数据。     

小木蠹蛾性行为和性信息素产生与释放的时辰节律

观察了小木蠹蛾Holcocerus insularis的性行为反应,并采用腺体提取、空气收集 、触角电位和田间试验等方法对雌蛾产生和释放性信息素的时辰节律进行了研究。结果表明： (1) 该虫羽化24 h后性成熟,婚飞和交配活动主要在1：00～4：00,交配历时15～45 min;(2) 大部分雌蛾一生交配1～3次,雄蛾多数一生只交配1次,雌雄比为1∶0.89; (3) 雌蛾腺体提取物中性信息素含量同蛾龄有关,2日龄雌蛾腺体性信息素含量最高;(4) 雌蛾腺体中性信息素含量在1：00时最高,而性信息素释放高峰在2：30。     

烟夜蛾雄蛾性附腺因子对雌蛾性信 息素合成的抑制作用

烟夜蛾Helicoverpa assulta处女蛾在交配后1 h,其性信息素滴度即显著降低,72 h内未见恢复。生测结果表明,烟夜蛾性信息素合成抑制因子主要来源于雄蛾性附腺。不同日龄雄蛾性附腺提取物的抑制活性无显著差异。光暗期对其活性具显著影响,暗期中雄蛾的性附腺物质对雌蛾性信息素合成具有较强抑制作用,而光期中雄蛾的性附腺物质不具抑制活性。在暗期的不同时间处理,对处女蛾性信息素合成的抑制作用无显著差异。雄蛾性附腺提取物对雌蛾性信息素合成的抑制作用与注射剂量有明显的相关性,0.2 ME（雄蛾当量）是产生显著抑制作用的最小剂量。对交配雌蛾注射性信息素生物合成激活神经肽（PBAN）提取物后,其性信息素合成又可恢复,这说明雌蛾交配后,性信息素滴度降低的原因是由于缺少了PBAN的调控。     

Electrophysiological and Behavioral Responses of Female Beet Armyworm Spodoptera exigua (Hübner) to the Conspecific Female Sex Pheromone

Electroantennogram (EAG) recordings showed that female Spodoptera exigua can detect their own sex pheromones (two single components and their mixture), displaying a similar dose–response pattern to that of males, although intensities of female responses were much less at all doses compared with males. Furthermore, the female calling behavior was inhibited and late-shifted by the presence of the female sex pheromone. When the pheromone components were presented, the calling female proportion in the peak calling period was significantly reduced and the calling peak time and calling termination time postponed, compared with controls. Although the calling behavior was inhibited, the pheromone titer of treated females was not different to the control, implying a reduced pheromone biosynthesis in the pheromone glands of treated moths. However, observations during the olfactometer experiments revealed that there were no obvious behavioral responses of females exposed to sex pheromone stimuli including whole gland extracts, 0.1, 1 or 10 μg binary pheromone mixtures.     

Diversity and function of male antennal glands in Cynipoidea (Hymenoptera)

The functional anatomy of antennal glands located either on the 3rd or on the 3rd and 4th antennomeres in males of several species of cynipoids was investigated. SEM observations revealed variously modified antennomeres with elevated plates, tyloids and excavated areas. In all the cases, the antennomeres are equipped with cuticular pores, corresponding internally to cuticular ducts. TEM studies showed the presence of type III integumentary glands, as classified by Noirot & Quennedey. Each glandular unit is made up of an innermost secretory cell, producing the secretion, and an outermost canal cell, producing the evacuating duct. The secretion passes through the duct and reaches the cuticular pores, concentrated in a ventro-lateral portion of the antennomere called the 'release and spread structure'. Both in Cynipidae and in Eucoilinae (Figitidae), the courtship behaviour includes a pre-copulatory phase characterized by intense antennal stroking. Bioassays in the eucoilins Leptopilina boulardi and L. heterotoma showed that these glands are the production site of a contact sex recognition pheromone, necessary for the female to accept the male.     

Female accessory reproductive gland activity in the yellow dung fly Scathophaga stercoraria (L.)

The role of the female accessory reproductive glands has been investigated in relatively few insects. Gland secretion has a number of potential functions, including lubrication during copula, involvement in fertilization and protection of eggs. Female yellow dung flies (Scathophaga stercoraria) have large paired accessory glands whose function(s) prior to this study were unknown. Our study indicated glands were involved in copulation and egg laying. The volume of secretion remaining in glands was negatively associated with copulation duration, and this effect was most pronounced in non-ovipositing females. Gland volume and secretion volume remaining in the glands were significantly smaller in females which were allowed to oviposit. In addition, there was a significant interaction between male size, female size and whether or not females were allowed to oviposit which affected the volume of the secretion remaining in the glands, with changes in secretion volume being greatest when males were large. Sperm were found in the accessory glands of some females and this was apparently not related to age, mating history of either sex, to female nutrition or male size. Our results indicate that either large males stimulate greater secretory responses from females or that females alter their responses based on male size.     

Location,morphology and histology of sex pheromone glands of the female gypsy moth, Lymantria dispar (L.)

Scanning electron microscopy, histology and a male wing fanning bioassay were used in this study to locate the sex pheromone-producing glands of the female gypsy moth, Lymantria dispar. When exposed to female sex pheromone, adult males exhibit a strong wing fanning behaviour prior to take off. We found that adult males showed positive response to calling females and to tissue extract from both dorsal and ventral portions of the intersegmental membrane between 8th and 9th-abdominal segments. A typical male response usually starts with elevation of antennae, movement of head in different directions, walking, wing fanning and onset of search flight. Histological and scanning electron microscopic studies suggested that the sex pheromone glands are located on the dorsal and ventral aspects of the intersegmental membrane. The glands appear as two highly convoluted integumentary areas with hypertrophied glandular epidermal cells.