Involvement of contact stimuli in the male-induced implantation block (the Bruce effect) in mice

Confinement of newly inseminated female mice exposed to confined alien males significantly decreased the male-induced implantation failure (the Bruce effect). By contrast, free (unconfined) females exposed to confined alien males showed a high rate of implantation failure. The results suggest that the pheromone of male origin involved in the Bruce effect is non-volatile and acts on the female through contact. The vomeronasal organ appears to be involved in the perception of this pheromone.     

Failure of familiar males to prevent alien male-induced implantation block (the Bruce effect) in laboratory mice

Implantation failure in newly inseminated females induced by exposure to alien males (the Bruce effect) was significantly reduced when the females were housed with the stud male. By contrast, newly inseminated females housed with a familiar male during exposure to alien males exhibited a high rate of implantation failure. The results suggest that the protective effect of the stud male on implantation is not because of the familiarity of the female with his odour cues. The results are consistent with the view that the newly inseminated female mouse identifies her coital partner as an individual because she becomes 'imprinted' with his odour during the pericopulatory period.     

Effect of cyproterone acetate on the pregnancy-blocking ability of male mice and the possible chemical nature of the pheromone

Even though castration abolished the ability of alien males to induce implantation failure (the Bruce effect) in newly inseminated females, treatment of intact alien males with the steroidal antiandrogen, cyproterone acetate, for 14 days (short term) did not significantly depress their ability to induce the Bruce effect. However, prolonged treatment (42 days) with cyproterone acetate suppressed the pregnancy-blocking ability of alien males to some extent, possibly due to the antigonadotrophic properties of the drug. Ovariectomized alien females treated with implants of testosterone (androgenized females) exhibited the ability to block implantation in newly inseminated females, but concurrent treatment of androgenized females with cyproterone acetate did not depress this ability. The results strongly suggest that the pheromone involved in the male-induced implantation failure is not the product of an androgen-dependent tissue, but is likely to be a product of androgen metabolism.     

Stud male-induced protection of implantation in food-deprived mice: masking effect of an artificial scent on pheromonal odour

In contrast to unscented stud males, stud males anointed with a commercial perfume failed to protect implantation in food-deprived females. It is suggested that the failure of perfumed stud males to protect pregnancy in their coital partners is due to the masking effect of the perfume on the stud male-originating olfactory cue which stimulates luteotrophic activity in females. The results are also consistent with the view that the newly inseminated female mouse identifies the stud male as an individual through a pheromonal cue and this is involved in the protective effect on implantation in the food-deprived female.     

Information asymmetries among males: implications for fertilization success in the thirteen-lined ground squirrel.

The classical view of mammalian mating competition focuses on combat and threat. By contrast, field studies have revealed that male mating success in some species is more strongly determined by mate location ability than by physical dominance. In thirteen-lined ground squirrels, competition in locating oestrous females is exacerbated by sperm competition that favours the first male to mate with a female. We used a female-removal experiment to identify the distinguishing characteristics of males that were the first at finding and mating with females. Each focal female was observed the day before she entered oestrus; the identities of males that made contact with her and the locations of their interactions were recorded. The following morning the females were temporarily removed, and we monitored male search behaviour in their absence. Males that arrived first were those that had spent more time with the female the previous day relative to their rivals. Time invested the day before, in turn, was highly correlated with male search persistence and familiarity with the female''s likely whereabouts. These results demonstrate that differential fertilization success can arise from information asymmetries among males: the advantaged individuals are those that have greater a priori knowledge of the reproductive state and spatial locations of prospective mates than rivals.     

Female mate choice across mating stages and between sequential mates in flour beetles

Few studies have examined how female premating choice correlates with the outcome of copulatory and post-copulatory processes. It has been shown that polyandrous Tribolium castaneum females discriminate among males before mating based on olfactory cues, and also exert cryptic choice during mating through several mechanisms. This study tested whether a male's relative attractiveness predicted his insemination success during copulation. Bioassays with male olfactory cues were used to rank two males as more and less attractive to females; each female was then mated to either her more attractive male followed by less attractive male, or vice versa. Dissections immediately after second copulations revealed a significantly higher percent of successful inseminations for females that remated with more attractive males compared with those that remated with less attractive males. These results indicate that cryptic female choice during copulation reinforces precopulatory female choice in T. castaneum, and suggest that females could use cryptic choice to trade up to more attractive males, possibly gaining better phenotypic or genetic quality of sires.     

Bromocriptine-induced implantation failure in mice: failure of the stud male to protect pregnancy.

The presence of the stud male failed to prevent implantation failure in newly inseminated females induced by administration of bromocriptine. This is in contrast with the ability of the stud male to prevent implantation failure in females induced by alien male exposure, and nutritional stress. Since bromocriptine is a potent inhibitor of hypophysial prolactin release by virtue of its stimulatory effect on hypothalamic dopaminergic activity, the results suggest that the stud male-originating luteotrophic stimulus is incapable of overriding the dopaminergic activity in bromocriptine-treated females.     

The Market Effect in the Wood Mouse, Apodemus sylvaticus:Selling Information on Reproductive Status

Grooming in the wood mouse is a means by which males obtain information about the reproductive state of females, as grooming creates a situation which allows the male to smell the groomed female’s anogenital area to ascertain her phase of oestrus. Although grooming is reciprocal in this species, it is asymmetrical in that males groom females more often than vice versa. This grooming asymmetry was studied using Markov chain analysis for grooming sequences in two captive wood mouse colonies, and transition rates were used to represent motivation in both sexes. Grooming sessions were often initiated by a male’s attempt to sniff an immobile female’s anogenital region, while the female would immediately react by avoiding or biting the male. In order to entice the female to remain, the male would begin grooming the female’s head and shoulder area, surreptitiously and consistently grooming downwards towards the female’s anogenital region, until she would again terminate such contact either by avoiding or biting the male. While, therefore, the male’s tendency to sniff the female’s anogenital region was stronger than his tendency to groom her, the female’s tendency to terminate the male’s naso-anal contact was much stronger than her tendency to terminate his grooming bouts. If the male did not initiate grooming after the female terminated naso-anal contact, she avoided further contacts and escaped. In mice, as in most mating systems, the demand for matings by males is far larger than the number of matings females offer. The mating market, therefore, is highly skewed, which gives females the opportunity to demand ‘commodities’ in return for allowing males to mate. This system allows females to ‘bargain’ with males to obtain grooming in return for anogenital contact. Females assess the length of time they receive grooming and will only allow males to attain anogenital contact after a certain threshold value has been passed. If anogenital contact provides the male with information about the female’s reproductive state and/or with sexual stimulation, then this process represents the first quantified example in short-lived mammals of females ‘selling sex’ in terms of the market effect. This paper therefore provides a new view of the regulation of grooming: grooming is not simply reciprocal with both participants concerned that the other does not ‘cheat’ (e.g. tit-for-tat (TFT)-like strategy), rather grooming is a commodity which can be bartered against female reproductive information or matings.     

Pregnancy blockage and the memory of the stud male in the vole (Microtus agrestis).

When female voles were allowed contact with the stud male for only 1 h at the time of mating, 55% exhibited pregnancy failure when exposed to a strange male 48 h later. When females were made psuedopregnant by hormone treatment and vaginal stimulation (i.e. no stud male involved), 87% exhibited luteal failure when exposed to a strange male. It is suggested that the characteristics of the stud male are rapidly imprinted upon the female at the time of mating and that this imprinting is important in preventing the female showing a blocking response to this male upon any subsequent exposure.     

Male housefly (Musca domestica L.) genital system as a source of mating pheromone

Male volatile mating pheromone is released by male houseflies placed in close proximity to females or in contact with female hydrocarbon extract. The pheromone induces the aggregation of virgin females in the vicinity of the male. Preparations of macerated testicular and ejaculatory duct tissue exert a similar effect, and these organs are, apparently, the source of the pheromone. Such preparations are not attractive to males or mated females. The pheromone also appears to induce female receptivity, since females that are exposed to emanations from male system preparations show a quicker rate of pairing with males. The pheromone effect results from the testes and ejaculatory ducts acting together, as preparations of each of these alone are inactive. Pheromone activity is age dependent and the greatest female reaction is observed when tissues of 6–10-day old males are used.     

Generalization of courtship learning in Drosophila is mediated by cis-vaccenyl acetate

Reproductive behavior in Drosophila has both stereotyped and plastic components that are driven by age- and sex-specific chemical cues. Males who unsuccessfully court virgin females subsequently avoid females that are of the same age as the trainer. In contrast, males trained with mature mated females associate volatile appetitive and aversive pheromonal cues and learn to suppress courtship of all females. Here we show that the volatile aversive pheromone that leads to generalized learning with mated females is (Z)-11-octadecenyl acetate (cis-vaccenyl acetate, cVA). cVA is a major component of the male cuticular hydrocarbon profile, but it is not found on virgin females. During copulation, cVA is transferred to the female in ejaculate along with sperm and peptides that decrease her sexual receptivity. When males sense cVA (either synthetic or from mated female or male extracts) in the context of female pheromone, they develop a generalized suppression of courtship. The effects of cVA on initial courtship of virgin females can be blocked by expression of tetanus toxin in Or65a, but not Or67d neurons, demonstrating that the aversive effects of this pheromone are mediated by a specific class of olfactory neuron. These findings suggest that transfer of cVA to females during mating may be part of the male's strategy to suppress reproduction by competing males.     

The roles of vision and olfaction in mate location by males of the tsetse fly Glossina morsitans morsitans

The roles of visual and/or olfactory stimuli in eliciting mating responses from male Glossina morsitans morsitans Westwood were examined, using a system for automatically recording the number and duration of mating strikes made towards decoys, under controlled conditions. The results confirm that there is no olfactory component of the female sex recognition pheromone sensed by the male antennae, and the attraction of males to females appears to be visual. The absence of male-male mating strikes was the result of the absence of female sex-pheromone, rather than the presence of a repellent mating deterrent in the male cuticle. Experiments with coloured, artificial, sex-pheromone-dosed, cotton decoys showed that colour had only weak effects on attractiveness and number of encounters with decoys, and that no colour caused significant enhancement of mating responses over those shown to decoy females.     

The Bruce effect in Norway rats

Intrauterine implantation of fertilized ova can be blocked by exposing recently inseminated females with an unfamiliar male. This selective pregnancy failure, designated as the Bruce effect (Bruce, Nature 1959; 184:105), is well studied in laboratory mice and has been confirmed in several other rodent species. However, no clear information exists concerning this phenomenon in the laboratory rat. The present study was conducted to investigate whether or not the Bruce effect exists in the rat. Females of two F1 hybrid strains (n(total) = 354) with different MHC genotypes (F344BNF1, RT1(lv1/n), and LEWPVGF1, RT1(l/c)) were mated with males of their own strain and subsequently exposed during the first 4 days postcoitus either to a male of the other hybrid strain or to an unfamiliar male of the same strain as the stud. The litter rate of each treatment group was determined. As a control, mated females of both strains were reexposed to the stud male to determine baseline litter rates. Female rats of both F1 hybrid strains showed a significantly lower litter rate when exposed to males of a different strain than their stud male, compared to the expected values of birth rates observed in control females (F344BNF1: P = 0.017; LEWPVGF1: P = 0.019). In contrast, there was no difference between expected and observed litter rates in females of both F1 hybrid strains after exposure to an unfamiliar male of the same strain as their stud. Our results demonstrate for the first time that the Bruce effect, well documented in mice, occurs in the Norway rat.     

Concurrent modulation of neuronal and behavioural olfactory responses to sex and host plant cues in a male moth

Mating has profound effects on animal physiology and behaviour, not only in females but also in males, which we show here for olfactory responses. In cotton leafworm moths, Spodoptera littoralis, odour-mediated attraction to sex pheromone and plant volatiles are modulated after mating, producing a behavioural response that matches the physiological condition of the male insect. Unmated males are attracted by upwind flight to sex pheromone released by calling females, as well as to volatiles of lilac flowers and green leaves of the host plant cotton, signalling adult food and mating sites, respectively. Mating temporarily abolishes male attraction to females and host plant odour, but does not diminish attraction to flowers. This behavioural modulation is correlated with a response modulation in the olfactory system, as shown by electro-physiological recordings from antennae and by functional imaging of the antennal lobe, using natural odours and synthetic compounds. An effect of mating on the olfactory responses to pheromone and cotton plant volatiles but not to lilac flowers indicates the presence of functionally independent neural circuits within the olfactory system. Our results indicate that these circuits interconnect and weigh perception of social and habitat odour signals to generate appropriate behavioural responses according to mating state.     

松墨天牛是否存在雌性接触信息素?

将松墨天牛MonochamusalternatusHope的身体不同部位切除之后的交配试验显示,分别剪去头部、鞘翅以及前胸部后仍然可以正常交配,用雄虫的腹部代替雌虫的腹部后,仍然能够引起雄虫的交配趋势动作,但不能完成真正交配,因此推测雌性接触信息素如果存在,主要集中在雌虫身体后半部,尤其是中后胸部位。但是冻死后的雌虫不再引起雄虫的交配行为,而且冻死的雌虫与活雌虫同时放入培养皿中,雄虫喜欢与活雌虫交配,不选择冻死的雌虫,因此推测视觉在其松墨天牛的交配过程中扮演着重要的角色,而接触性信息素有可能并不存在。将雄虫的触角分别从第5~6节和第1节剪去的试验结果显示,雄虫仍然可以很快地找到雌虫,发生交配行为。这有2种可能的解释一是雄虫不仅仅靠触角来感受雌虫的接触信息素,头部下颚须和下唇须上或者足上也有可以感受信息素的感受器;二是雌虫不存在接触信息素,雄虫通过视觉就可以找到对方,完成交配过程。浸泡致死的雌虫不再引起雄虫的交配行为,并且将正己烷和乙醚的浓缩液滴加在浸泡致死的雌虫以及玻璃棒和鹅卵石的生测试验结果显示,都没有引起雄虫的交配行为。这进一步说明,松墨天牛可能并不存在体表接触信息素。性成熟后的松墨天牛雌雄成虫是靠衰弱寄主的强烈的挥发性气味聚集到寄主树干上,在树干这个较为狭小的环境里,有可能是通过视觉找到配偶,完成交配过程。     

Induction of dominant lethals in progeny of male CBA mice after pheromonal action

The inhibiting effect of pheromone 2,5-dimethylpyrazine of house mouse females on the reproductive function of the CBA male mice was studied. The mutagenic effect of six-day pheromonal effect was assessed by dominant lethal test. Analysis for the frequency of dominant lethals showed that the pheromonal effect results in an increased death rate of the progeny of the treated males. This is probably explained by implantation failure and is expressed in a reduced average number of the implantation sites and low live embryos per female. The proportion of females with live embryos decreased significantly. The implication of the effect of female mouse pheromone 2,5-dimethylpyrazine on the genetic processes in germ cells of male mice is discussed.     

Long-term effects of accelerated or delayed sexual maturation on reproductive output in wild female house mice (Mus musculus)

The effects of acceleration and delay of puberty in female house mice on survival and reproduction were tested using 6 experimental groups: (1) control females mated at the time of first oestrus, (2) females mated at weaning, (3) females treated with male urine starting at weaning and mated at first oestrus, (4) females housed in groups and mated at first oestrus, (5) females housed alone, treated with urine from grouped females and mated at first oestrus, and (6) females housed alone and mated at 68 days of age. Females caged with males at weaning or treated with male urine and mated at puberty had lower rates of survival to 180 days of age, but did not differ in rates of fertility from mice in the other four treatments. Those females that were housed with males from weaning or treated with male urine also had smaller total numbers of litters, fewer total young, and smaller average litter sizes than did females for which the age of mating was delayed, by grouping or treatment with urine from grouped females, or by being held until age 68 days before mating. Control females mated at first oestrus generally were intermediate or did not differ from the male treatments on these dependent variables. There were no differences in the average number of female young/litter across the 6 treatments. However, females that were delayed in age of first mating had significantly more male young/litter than did females that were accelerated in their sexual development or control females.(ABSTRACT TRUNCATED AT 250 WORDS)     

Love Is Blind: Indiscriminate Female Mating Responses to Male Courtship Pheromones in Newts (Salamandridae)

Internal fertilization without copulation or prolonged physical contact is a rare reproductive mode among vertebrates. In many newts (Salamandridae), the male deposits a spermatophore on the substrate in the water, which the female subsequently takes up with her cloaca. Because such an insemination requires intense coordination of both sexes, male newts have evolved a courtship display, essentially consisting of sending pheromones under water by tail-fanning towards their potential partner. Behavioral experiments until now mostly focused on an attractant function, i.e. showing that olfactory cues are able to bring both sexes together. However, since males start their display only after an initial contact phase, courtship pheromones are expected to have an alternative function. Here we developed a series of intraspecific and interspecific two-female experiments with alpine newt (Ichthyosaura alpestris) and palmate newt (Lissotriton helveticus) females, comparing behavior in male courtship water and control water. We show that male olfactory cues emitted during tail-fanning are pheromones that can induce all typical features of natural female mating behavior. Interestingly, females exposed to male pheromones of their own species show indiscriminate mating responses to conspecific and heterospecific females, indicating that visual cues are subordinate to olfactory cues during courtship.     

Pheromone detection by a pheromone emitter: a small sex pheromone-specific processing system in the female American cockroach

Many animals depend on pheromone communication for successful mating. Sex pheromone in insects is usually released by females to attract males. In American cockroaches, the largest glomerulus (B-glomerulus) in the male antennal lobe (first-order olfactory center) processes the major component of sex pheromone. Using intracellular recordings combined with fine neuroanatomical techniques, we provide evidence that the female homolog of the male B-glomerulus also acts as a sex pheromone-specific detector. Whereas ordinary glomeruli that process normal environmental odors are innervated by single projection neurons (PNs), the B-glomerulus in both sexes is innervated by multiple PNs, one of which possesses a thicker axon, termed here B-PN. Both soma size and axon diameter were smaller on B-PNs from females compared with B-PNs from males. The female B-PNs also produce fewer terminal arborizations in the protocerebrum than male B-PNs. Termination fields in the lateral protocerebrum of the female B-PN are mostly segregated from those formed by other uniglomerular PNs innervating ordinary glomeruli. Female B-PN activity was greatest in response to sex pheromone but lower than that in the male B-PN. This specific detection system suggests that sex pheromone affects the behavior and/or endocrine system of female cockroaches.     

Effects of sex pheromone in electrostatic powder on mating behaviour by Lobesia botrana males

Abstract:  In laboratory trials, mating behaviour of males of Lobesia botrana Den. and Schiff. (Lep., Tortricidae) was investigated after treatment with an electrostatically chargeable powder, Entostat^TM. Male moths were powdered with either blank Entostat or Entostat containing sex pheromone. Significantly more Entostat adhered to L. botrana males when sex pheromone was added to the powder compared with blank Entostat. Powdering male moths with Entostat, with or without sex pheromone, caused a significant reduction in antennal response when antennae were placed 2 cm away in downwind direction, while there was little effect when antennae were placed 25–100 cm from the EAG in downwind direction. In a flight study, powdering males of L. botrana caused significant increase in time before flight initiation and reduction in proportion of males making contact with calling females compared with untreated males. In a mating experiment in Petri dishes (to minimize required search time to locate female), significantly fewer females mated successfully (based on dissection of bursa copulatrix) when males had been treated with pheromone-loaded powder. Overall, powdering males of L. botrana caused considerable suppression of mating behaviour on various levels, and these suppressing effects were increased after adding sex pheromone to Entostat.