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Behavioral responses of sexually active mud snails: kariomones and pheromones.

We postulated that kariomones and pheromones function to organize egg capsule deposition and breeding aggregations of mud snails (Ilyanassa obsoleta). Sexually active male and female mud snails were exposed to water-born odors from juvenile snails, nonaggregated female snails, oysters, egg capsule depositing snails, sexually active males, sexually active females, and copulating snails and tested for a follow response using a ring assay. Neither sex responded to juvenile odor or to odor of sexually inactive females. Snails turned and followed water born odors from oysters, when tested with the odors of sexually active males, sexually active females, and copulating pairs, we found sex-specific differences in responses. Sexually active females followed the odor of sexually active males and ignored sexually active female odor. Sexually active males followed the odor of sexually active females and ignored sexually active male odor. Both sexes followed the odor of copulating pairs and egg capsule depositing snails. Snail activities in breeding aggregations are organized by at least three pheromones and one kariomone, and reflect the multifunctionality of the aggregations.

Animals↗

Differential effects of yohimbine and naloxone on copulatory behaviors of male rats.

Prior research has demonstrated that both yohimbine, an alpha2-adrenergic antagonist, and naloxone, an opiate antagonist, facilitate components of copulatory behaviors in nonstressed male rats. In the present experiments, we demonstrate that these drugs differentially affect copulatory behaviors when the behavioral testing situation contained an aversive element. Male rats received an injection of lithium chloride (0.3 M, 20 ml/kg, ip) immediately after each encounter with an estrous female. Consequently, male copulatory behaviors gradually declined during successive test sessions. These male rats also received ip injections of either yohimbine (2 mg/kg/ml), naloxone (4 mg/kg/ml), or isotonic saline 20 min prior to each copulation test. Yohimbine-treated rats were more likely to copulate than control rats during both acquisition and extinction of lithium chloride-induced associative inhibition of copulatory behavior. Conversely, naloxone-treated rats were less likely to copulate than control rats during both acquisition and extinction. These data are consistent with the hypothesis that yohimbine increases sexual motivation in the male rat and limit the generality of the excitatory effects of naloxone on copulatory behaviors.

Animals↗

Levels of serum steroids, aromatase activity, and estrogen receptors in preoptic area, hypothalamus, and amygdala of B6D2F1 male house mice that differ in the display of copulatory behavior after castration.

Most male B6D2F1 hybrid house mice continue to copulate after castration (continuers), whereas others do not (noncontinuers). Copulation in continuers appears estrogen dependent. Serum testosterone (T), 17 beta-estradiol (E2), and dihydrotestosterone (DHT), as well as aromatase activity (AA) and estrogen receptor (ER) levels in preoptic area (POA), hypothalamus (HYP), and amygdala (AM) were measured to determine if continuers and noncontinuers differ in estrogen physiology. In general, continuers and noncontinuers did not differ in serum steroid levels, AA, or ER levels. Castration reduced AA in the POA, HYP, and AM. Castration did not affect nuclear ER levels in the POA and HYP but reduced nuclear ER in AM. The data demonstrate that castrated B6D2F1 male mice continue to be under the influence of circulating nongonadal E2 that is important for copulation.

Amygdala↗

Impaired sexual response after lesions of the paraventricular nucleus of the hypothalamus in male rats.

N-methyl-D-aspartic acid (NMDA) or radiofrequency (RF) lesions were made in the paraventricular nucleus of the hypothalamus (PVH) of male rats. The rats were tested for copulation, noncontact erection (NCE) evoked by remote cues from estrous females, and (after RF lesions) reflexive erection. NMDA, which destroyed parvocellular but spared magnocellular neurons, caused no deficits in copulation but caused longer NCE latencies and fewer NCEs. Rats with RF lesions had parvo- and magnocellular neuron damage; these males copulated to ejaculation, but they had lower intromission ratios and longer ejaculatory latencies. RF-lesioned rats also had longer NCE latencies, and a smaller proportion of males displayed reflexive erection. Results indicate that the PVH participates in mediating erectile function in copula and ex copula, and that the parvo- and magnocellular PVH neurons may have different roles in mediating erection.

Animals↗

Sexual fetishism in a quail (Coturnix japonica) model system: test of reproductive success.

In the present study, the authors explored the reproductive consequences of fetishistic behavior in a previously developed animal model of sexual fetishism (F. Köksal et al., 2004). Male domesticated quail (Coturnix japonica) received sexual conditioning trials in which a terrycloth object (the conditioned stimulus [CS]) was paired with the opportunity to copulate with a female quail (the unconditioned stimulus). Approximately half of the male quail came to copulate with the CS object and were considered to have developed fetishistic behavior. Each of the male quail was then tested with a female quail, whose eggs were incubated to determine rates of fertilization. The CS object was present for 30 s before and during the copulation test. Fetishistic male quail were slower to achieve cloacal contact with the female quail and showed less efficient copulatory behavior. However, they fertilized a greater proportion of eggs than nonfetishistic male quail. These results are unexpected from previous studies of the relationship between reproductive success and copulatory behavior and are discussed in terms of how fetishistic behavior directed toward an inanimate object may modify male-female interactions.

Animals↗

Regulation of female sexual behavior in the golden hamster: behavioral effects of mating and ovarian hormones.

In the female hamster, sexual receptivity was abbreviated following copulation. Short-term effects of mating, lasting approximately 24-48 hr, were observed in females mated in either estradiol- or estradiol-and-progesterone-induced estrus. Long-term effects of copulation, of 9 days or more in duration, were apparent only in females chronically treated with both estradiol and progesterone, suggesting that progesterone exposure prolongs the inhibitory effects of mating. When progesterone stimulation was intermittent through the use of short-acting injection procedures, recovery from mating could be detected in 48 hr and was complete within 96 hr after copulation. The pattern of behavioral response to ovarian hormones in unmated females was also documented, replicating the "biphasic" effect of progesterone in this species. The inhibitory effects of mating and progesterone summate to produce consistent decrements in female sexual receptivity. It is postulated that short-term postcopulatory abbreviations in receptivity would reduce the vulnerability of the estrous female while long-term inhibitions, interacting with sustained progesterone stimulation, would reduce the probability of mating during pregnancy when hormone levels are elevated for a prolonged period.

Adrenalectomy↗

No evidence for illegitimate young in monogamous and polygynous warblers.

In animals with internal fertilization, paternity is uncertain. In birds, the occurrence of copulations outside the pair-bond has been documented in a number of species, but the extent to which these result in illegitimate young is largely unknown, and constitutes a major deficiency in our understanding of avian mating systems. The analysis of tandemly repeated sequences (minisatellites), has enhanced our ability to make individual identifications and paternity determinations. Here we describe the use of a bird minisatellite DNA probe in assigning paternity in natural populations of the monogamous willow warbler Phylloscopus trochilus and of the polygynous wood warbler Phylloscopus sibilatrix. In both species this probe detects a multiple locus pattern and a single locus that exhibits a variable number of tandem repeats. Although we observed intrusions by non-resident males into the territories of paired males and extra-pair copulations, no illegitimate offspring were detected among 176 young from 32 families of both species, implying that extra-pair copulations have little or no genetic impact.

Alleles↗

Female feral fowl eject sperm of subdominant males.

Paternity is often determined by competition between the ejaculates of different males. Males can also use particular behaviours or structures to manipulate how females use sperm. However, the ability of females to bias sperm utilization in favour of preferred males independently of male manipulation has not been demonstrated. Females are predicted to respond differentially to the sperm of different males when the reproductive interests of the sexes differ and when females are coerced into copulating. Here we show that in female feral fowl most copulations are coerced, and that females consistently bias sperm retention in favour of the preferred male phenotype. Females prefer to copulate with dominant males, but when sexually coerced by subordinate males, they manipulate the behaviour of dominant males to reduce the likelihood of insemination. If this fails, females differentially eject ejaculates according to male status in the absence of any male manipulation and preferentially retain the sperm of dominant males.

Animals↗

Courtship Disruptions and Male Mating Strategies: Examples from Female-Defense Mating Systems.

Males frequently interrupt the copulation attempts of other males, and these courtship disruptions may limit the extent to which a few males are able to monopolize mating access to females. Males actively defend sexually receptive females in many species in which females form dense aggregations during the breeding season. Across and within such species there is considerable variation in the mating tactics adopted by males, with males in some cases defending groups of females and in other cases sequentially consorting with individual females. Colonial blackbirds have been central to studying this mating system, and we develop a conceptual model for how courtship disruption may account for variation in male mating tactics in this group. Our model assumes that the frequency of disruptions increases with greater colony size. As a consequence, successful copulations are less likely to occur at large colonies than at small colonies, and males are expected to switch from defending multiple females at the colony to consorting individual females away from it. Results from two species of blackbird support the basic assumptions of this model. In one species, the Montezuma oropendola, disruptions occur rarely and males defend groups of females, whereas in the other species, the yellow-rumped cacique, disruptions are frequent and males defend single females. Moreover, consistent with a key prediction, within each species, males associated with small colonies remain at the colony and defend groups of females, whereas males spend little time defending groups of females at large colonies and rarely attempt copulations there. This model has the potential to explain variation in male mating strategies and female monopolization for other taxa in which females form breeding aggregations.

Icteridae↗

Sexual behaviour during ovarian cycles, pregnancy and lactation in group-living stumptail macaques (Macaca arctoides).

Sexual interactions were studied in a large, well-integrated group of stumptail macaques, continuously living together. Copulations with the highest-ranking male and with young males were equally distributed over the ovarian cycle. Copulations with other adults males also occurred during all cycle phases, but were most frequent peri-ovulatory. This was most distinct in conceptive cycles. Copulatory activity continued during early pregnancy, but was virtually absent during the middle and latter part of pregnancy and during lactation. Fluctuations in copulation frequency could be attributed to fluctuations in female attractivity (as measured by directed male masturbations), but not in proceptivity (spontaneous presents) or receptivity (acceptance ratios). However, hormonal influences on attractivity could be overruled by social factors. With regard to the expression of heterosexual behaviour, stumptail macaques seem to be less hormone-dependent than other primate species studied, both in the laboratory and under more natural circumstances.

Animals↗

Ovulation and fertilization in the vole, Pitymys subterraneus.

The females of Pitymys subterraneus bred in laboratory conditions have an irregular sexual cycle and induced ovulation. The first freshly ovulated eggs, surrounded by dense cumulus cells, appear in oviducts 10 h after copulation. Administering exogenous gonadotropins: pregnant mare's serum (PMS), human chorionic gonadotropin (hCG) or luteinizing hormone releasing hormone (LHRH), also induces ovulation in mature females of Pitymys subterraneus. In these experimental conditions females ovulate a similar number of eggs as after copulation. Dual stimulation (PMS and hCG plus copulation) does not result in the ovulation of a large number of normal ova, however, it does cause a release of degenerated oocytes.

Animals↗

Luteinizing hormone and prolactin in mated female meadow voles housed in long and short day lengths.

Fertility differs dramatically between female meadow voles (Microtus pennsylvanicus) housed in long (LD; 14 h light/day) and short (SD; 10 h light/day) day lengths. All mated LD females ovulate, and 75-100% produce litters. In contrast, 40% of SD females that mate only after long contact with a male do not ovulate, and fertility for SD females is much lower (30-40% produce litters overall). Because copulation causes the surge in LH required for ovulation and the increase in prolactin (PRL) needed to maintain corpora lutea, we hypothesized that SD females might have lower ovulation rates and litter production because of inadequate hormonal responses. Serum LH was measured 24 h before, and 30, 60, 90, 120, and 180 min after, mating. Prolactin was measured before, and 24 and 48 h after, copulation. LD and SD females that ovulated demonstrated a sharp increase in LH levels at 30 and 60 min post mating, but LH in non-ovulating females was approximately 50% lower than that of ovulating females at 30 and 60 min post mating. PRL in LD females was double baseline levels at 24 and 48 h after mating. PRL in SD females increased at 24 h, but declined to pre-mating levels by 48 h post mating. These data suggest that reduced fertility in short day lengths in female voles may be related to two separate problems: 1) The rise in LH following copulation is insufficient in some females to cause ovulation. 2) PRL may not remain elevated long enough to support functional corpora lutea in some females that ovulate.

Animals↗

Effect of repeated collection on semen characteristics of alpacas.

Semen characteristics of alpacas were studied after repeated collections. Twelve adult males were divided into three groups of four each for semen collection once, twice, or three times every other day. The duration of copulation; volume of ejaculate; pH; motility; sperm concentration (number of sperm/milliliter semen); total number of sperm per ejaculate; and percentages of live, normal, and abnormal spermatozoa were analyzed by regression analysis. Semen color and consistency were analyzed by the chi-square test. Between the first, second, and third ejaculations, there were differences (p < 0.05) in sperm concentration; percentages of normal spermatozoa and abnormal spermatozoa; sperm with abnormal heads and abnormal tails; and consistency (viscous, viscous, and semi-viscous). There were no differences (p > 0.05) in ejaculated volume, percentage of live spermatozoa, pH, percentage of cytoplasmic droplets, and duration of copulation. Some males from which semen was collected on the three-mating schedule ejaculated only seminal plasma during the second and third copulation starting on Day 10 of the study. There were differences between males (p < 0.05) for most of the characteristics studied. In sum, frequency of mating affected some semen characteristics that may be important determinants of the fertility of male alpacas.

Animals↗

Death feigning in the face of sexual cannibalism.

Pre-copulatory sexual cannibalism by females affects male and female reproductive success in profoundly different ways, with the females benefiting from a meal and the male facing the risk of not reproducing at all. This sexual conflict predicts evolution of traits to avoid cannibalism and ensure male reproductive success. We show that males of the nuptial gift-giving spider Pisaura mirabilis display a remarkable death feigning behaviour--thanatosis--as part of the courtship prior to mating with potentially cannibalistic females. Thanatosis is a widespread anti-predator strategy; however, it is exceptional in the context of sexual selection. When the female approached a gift-displaying male, she usually showed interest in the gift but would sometimes attack the male, and at this potentially dangerous moment the male could 'drop dead'. When entering thanatosis, the male would collapse and remain completely motionless while retaining hold of the gift so it was held simultaneously by both mates. When the female initiated consumption of the gift, the male cautiously 'came to life' and initiated copulation. Death feigning males were more successful in gaining copulations, but did not have prolonged copulations. We propose that death feigning evolved as an adaptive male mating strategy in conjunction with nuptial gift giving under the risk of being victimized by females.

Animals↗

Mate guarding in the Seychelles warbler is energetically costly and adjusted to paternity risk.

Males may increase their fitness through extra-pair copulations (copulations outside the pair bond) that result in extra-pair fertilizations, but also risk lost paternity when they leave their own mate unguarded. The fitness costs of cuckoldry for Seychelles warblers (Acrocephalus sechellensis) are considerable because warblers have a single-egg clutch and, given the short breeding season, no time for a successful replacement clutch. Neighbouring males are the primary threat to a male's genetic paternity. Males minimize their loss of paternity by guarding their mates to prevent them from having extra-pair copulations during their fertile period. Here, I provide experimental evidence that mate-guarding behaviour is energetically costly and that the expression of this trade-off is adjusted to paternity risk (local male density). Free-living males that were induced to reduce mate guarding spent significantly more time foraging and gained significantly better body condition than control males. The larger the reduction in mate guarding, the more pronounced was the increase in foraging and body condition (accounting for food availability). An experimental increase in paternity risk resulted in an increase in mate-guarding intensity and a decrease in foraging and body condition, and vice versa. This is examined using both cross-sectional and longitudinal data. This study on the Seychelles warbler offers experimental evidence that mate guarding is energetically costly and adjusted to paternity risk.

Animals↗

How well do mating frequency and duration predict paternity success in the polygynandrous water strider Aquarius remigis?

The relationship between mating success and paternity success is a key component of sexual selection but has seldom been estimated for species in which both sexes mate with many partners (polygynandry). We used a modification of Parker's sterile male technique to measure this relationship for the water strider Aquarius remigis in 47 laboratory populations simulating natural conditions of polygynandry. We also tested the hypothesis that prolonged copulation, a characteristic of this species, enhances paternity success. Mating behavior and paternity success were assayed for four days while males and females freely interacted. Paternity success was also assayed for an additional 7 days when females were isolated from males. Mating success significantly predicted paternity success and accounted for < or = 36% of the variance. Copulation duration was negatively related to both mating success and paternity success and did not explain any of the residual variance in paternity success. Thus, we found no evidence that prolonged copulation functions as a paternity assurance strategy in this species. Comparisons of sterile and fertile males suggested that paternity success is directly influenced by the quantity of sperm transferred. Our results support previous studies that have used mating success to estimate sexual selection, but also highlight the potential importance of sperm competition and other postinsemination processes.

Animals↗

Fecundity and MHC affects ejaculation tactics and paternity bias in sand lizards.

We demonstrate that extending copulation enhances probability of paternity in sand lizards and that determinants of copulation duration depend on a males' mating order (first or second). First males, with no information on presence of rivals, extend copulation when mating with a more fecund female. Second males, however, adjust copula duration in relation to a first male's relatedness with his female, which there is reason to believe can be deduced from the MHC-related odor of the copulatory plug. Male-female relatedness negatively influences a male's probability of paternity, and when second males are in a favored role (i.e., the first male is the one more closely related to the female), second males transfer larger ejaculates, resulting in higher probability of paternity. This result corroborates predictions from recent theoretical models on sperm expenditure theory incorporating cryptic female choice and sexual conflict. More specifically, the results conform to a "random roles" model, which depicts males as being favored by some females and disfavored by others, but not to a "constant-type" model, in which a male is either favored or disfavored uniformly by all females in a population.

Animals↗

Opposing roles of the nucleus accumbens and anterior lateral hypothalamic area in the control of sexual behaviour in the male rat.

Opposing roles have been implicated for the nucleus accumbens (NAc) and anterior portion of the lateral hypothalamic area (aLHA) in the regulation of sexual behaviour in male rats based on in vivo neurochemical correlates. The present study provides functional evidence supporting this hypothesis by examining the effects of lesions to these structures on copulation, noncontact erection and receptive female preference. Sexually naïve male Long-Evans rats received either bilateral 1.0- micro L injections of NMDA (10 micro g/ micro L/side) or vehicle (shams) into either the aLHA or the NAc. During repeated tests of copulation most of the sham-lesioned males, but few of the aLHA-lesioned and NAc-lesioned males, copulated to ejaculation. Most of the NAc-lesioned males also failed to intromit, whereas the majority of the aLHA-lesioned males intromitted repeatedly. During exposure to an inaccessible receptive female behind a wire-mesh screen, aLHA-lesioned males displayed facilitation of noncontact erections, whereas NAc-lesioned males displayed impaired noncontact erections. Conversely, during simultaneous exposure to inaccessible receptive and nonreceptive females in different compartments, all males spent more time in the proximity of the receptive female. These findings indicate that the aLHA plays an inhibitory role in the regulation of sexual arousal and an excitatory role in the regulation of ejaculation. Conversely, the NAc plays an excitatory role in the regulation in sexual arousal.

Animals↗