Mate choice and sexual conflict in <Emphasis Type="Italic">Brachionus plicatilis</Emphasis> (Rotifera)

Male mate choice in Brachionus plicatilis is based on information from coronal receptors and contact chemoreception of a glycoprotein signal on the body surface of females. Male mating behavior is modulated by mating signal quality and strength, which determines male mating persistence and circling intensity. We probe the sensory abilities of males by better resolving their discrimination of female age. Males preferentially initiate mating with young females, but not too young. Males circle 0.2 h old females just hatched from resting eggs only about 1/2 as frequently as 12 h old females. Males are even more discriminating of females with whom they copulate, preferring 3 h old females significantly more than 0.2 h or 6 h old females. In contrast, males cannot distinguish between virgin females and those who have already copulated. Similarly, males circled and copulated with females hatching from resting eggs with equal frequency as those hatching from amictic eggs. The counterpoint to male mate choice is female resistance to mating. When males contact females, the females respond with one of four behaviors: no response, acceleration, foot flipping, or coronal retraction. In 65% of male–female encounters, there was no initial response by females. However, when males began circling females, females accelerated 11.1 times more often than when males were absent. The second type of evasive female behavior was foot-flipping, which tends to knock off males attempting to circle the female. In the presence of circling males, females performed foot-flipping behavior three times more often than in the absence of males. Coronal retraction, where they stop swimming and withdraw their corona, was observed less frequently than acceleration or foot-flipping, and there was no difference in the presence or absence of males. These data are interpreted in the context of sexual conflict, where the behaviors that optimize male and female fitness differ. Guest editors: S. S. S. Sarma, R. D. Gulati, R. L. Wallace, S. Nandini, H. J. Dumont & R. Rico-Martínez Advances in Rotifer Research     

Multiple mechanisms of phenotype development in the bluehead wrasse

Despite having detailed information on mechanisms mediating sex-typical behavior in many species, we have little understanding of whether the same mechanisms regulate these behaviors when they are performed in the same species under different social contexts. In the five field experiments of this study of bluehead wrasses (Thalassoma bifasciatum), a sex-changing fish, we examined the roles of arginine vasotocin (AVT) and the potent teleost androgen 11-ketotestosterone (11KT) in mediating sexual and aggressive behaviors typical of dominant males. We demonstrated that AVT appears necessary for the assumption of dominant territorial status in males and females, but is sufficient only in the socially dominant terminal phase (TP) male phenotype. Specifically, an AVP V(1) receptor antagonist prevented both TP males and females from gaining dominance over recently vacated territories. However, unlike TP males in a previous study, neither females nor initial phase males responded to AVT treatment with increases in display of TP male typical behaviors when under social conditions that inhibit sex change. Treating females with 11KT did not alter responsiveness to AVT, but did induce male coloration and courtship behavior that was not observed in oil-treated females. Combined with the results of a previous study, these results indicate that the ability of AVT to induce male-typical behavior differs among sexual phenotypes and that this differential responsiveness appears to be dependent on social context and not directly on exposure to 11KT. Furthermore, since 11KT can induce courtship behavior in females that is not affected by AVT, there may be different hormonal mechanisms mediating courtship behavior under different social contexts.     

Discrimination of conspecific individuals via cuticular pheromones by males of the cricket Gryllus bimaculatus

Cuticular substances on the body surface of crickets serve as pheromones that elicit a variety of different behaviors in male crickets. Antennal contact between males and females resulted in courtship behavior, and that between two males resulted in aggressive displays. As a first step in elucidating how crickets recognize and discriminate individuals, behavioral responses of male individuals to cuticular substances of conspecific males or females were investigated. The behavioral responses of males to antennal or palpal stimulation with an isolated antenna from a male or a female were recorded. To both antennal and palpal stimulation with female antennae, the majority of males responded with courtship behavior; to stimulation with male antennae, males responded with aggressive displays. To gain insight into the chemical nature of the behaviorally relevant components, isolated antennae were washed in either n-hexane, acetone or ethanol before behavior assays. Washed antennae no longer elicited courtship or aggressive responses in males. Next, polypropylene fibers were smeared with substances from the body surface of females and used for antennal stimulation. This experiment showed that the quality and quantity of cuticular substances appear to be highly age-dependent. Significantly more males responded with courtship behavior to cuticular substances from younger females. Isolated males generally showed higher levels of aggression than males reared in groups. Grouped males also were more likely to display courtship behavior towards antennae from younger females, and aggressive behavior towards antennae from older females. These results suggest that male discrimination of mating partners depends on the nature of female cuticular substances.     

Exploring the influence of individual courtship behaviors on male mating success in a blow fly

Male courtship behavior and displays influence female mating decisions, and therefore affect mating success in a diverse range of organisms. While there is substantial evidence confirming that females prefer males who invest more in courtship, less attention has been paid to the relative importance of individual behaviors, and the discrete sequences of courtship that result in mating success. The small hairy maggot blow fly, Chrysomya varipes, performs stereotyped courtship behaviors, involving orienting, tapping, waving, arching, wing vibration and mounting. This study aimed to quantify male investment in specific courtship behaviors, and compare courtship investment and behavioral transitions between males who gained mating success (successful males) and those that did not (unsuccessful males). Our results show that mating success was influenced by the behaviors orienting, tapping, arching and mounting. Behavioral transitions revealed a distinct pattern of behaviors leading to a mating attempt, and some differences were observed between successful and unsuccessful males. Overall, our findings suggest that female mating decisions were based on differences in specific male courtship behaviors. This detailed observational study has quantified multiple courtship behaviors for the first time in C. varipes, and highlights the importance of considering multiple behavioral traits when exploring the influence of male courtship on mating success.     

雌星豹蛛性信息素的行为学证据

采用星豹蛛(Pardosa astrigera)成熟雄蛛求偶时潜伏时间、静止时间、身体震动和第一墩步足伸展次数等行为参数,利用行为学方法测定了不同性别、日龄和生殖状态的星豹蛛雌蛛释放的拖丝对雄蛛求偶行为的影响。结果表明,雄蛛第一对步足伸展和身体震动等典型求偶行为是进行星豹蛛性信息素生物测定的可靠评价指标。星豹蛛雄蛛能通过拖丝上的性信息素辨别星豹蛛的性别、日龄和生殖状态。雄蛛在成熟3周未交配雌蛛拖丝处理过滤纸上潜伏时间和静止时间都相应最短,在交配未产卵雌蛛、雌亚成蛛和成熟雄蛛拖丝上时间中等,在卵孵化雌蛛拖丝处理滤纸上潜伏时间和静止时间都相应最长。成熟3周未交配雌蛛和交配未产卵雌蛛释放的拖丝都能引起雄蛛第一对步足伸展和身体震动等典型求偶行为,雄蛛对成熟3周未交配雌蛛拖丝典型求偶行为的频率都相应高于交配未产卵雌蛛。卵孵化雌蛛释放的拖丝虽也能引起雄蛛第一对步足伸展行为,但其伸展频率显著降低;而其它拖丝都不能引起雄蛛典型求偶行为。     

Complex Courtship Behavior in the Striped Ground Cricket, Allonemobius socius (Orthoptera: Gryllidae): Does Social Environment Affect Male and Female Behavior?

Complex courtship in the striped ground cricket, Allonemobius socius, involves a series of behaviors alternating between the sexes. We examined if complex courtship allows either or both genders to evaluate their mate and how mating behavior changes in different social environments. While complex courtship may allow discrimination by both sexes, here only females exhibited a preference. Males did not alter their courtship behavior or change spermatophore size for different size females. In contrast, females initiated copulation more quickly with bigger males possessing bigger spermatophores. In a different social environment (additional male, female, or both), males were less likely to omit courtship songs and female discrimination of mates changed. The distinct differences in male and female behavior suggest that subtle changes in social environment can have important consequences in structuring courtship and mating behavior.     

fruitless splicing specifies male courtship behavior in Drosophila

All animals exhibit innate behaviors that are specified during their development. Drosophila melanogaster males (but not females) perform an elaborate and innate courtship ritual directed toward females (but not males). Male courtship requires products of the fruitless (fru) gene, which is spliced differently in males and females. We have generated alleles of fru that are constitutively spliced in either the male or the female mode. We show that male splicing is essential for male courtship behavior and sexual orientation. More importantly, male splicing is also sufficient to generate male behavior in otherwise normal females. These females direct their courtship toward other females (or males engineered to produce female pheromones). The splicing of a single neuronal gene thus specifies essentially all aspects of a complex innate behavior.     

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.     

Analysis of the Courtship Behavior of the Navel Orangeworm, Amyelois transitella (Walker) (Lepidoptera: Pyralidae), with a Commentary on Methods for the Analysis of Sequences of Behavioral Transitions

The courtship behavior of the navel orangeworm, Amyelois transitella, was examined in a wind tunnel. Sixty nine courtship sequences were analyzed and successful sequences divided into two categories: rapid courtship sequences, which involved few breaks in contact, short or no periods of male/female chasing and lasted <10 s between initial contact and mating; and prolonged courtship sequences, which involved many breaks in contact, extended periods of male/female chasing and lasted >10 s. Fifty six (81%) courtships were successful (50.7% rapid courtship and 30.4% prolonged courtship); the remaining 13 (18.8%) sequences were failed courtships. Of failed courtships, 9 (13.0%) were due to males losing contact with females during courtship chases and 4 (5.8%) due to females flying away immediately after male contact. Of all courtship sequences involving a break in contact during a chase, 38.5% resulted in an unsuccessful mating attempt. These findings contrast with previous studies of the courtship behavior of the navel orangeworm, potentially indicating that the type of bioassay used to study courtship may have a large effect on the behavioral sequences displayed. We evaluate several diagnostic techniques for the analysis of sequences of behavioral transitions.     

Social context influences aggressive and courtship behavior in a cichlid fish

Social interactions require knowledge of the environment and status of others, which can be acquired indirectly by observing the behavior of others. When being observed, animals can also alter their signals based on who is watching. Here we observed how male cichlid fish (Astatotilapia burtoni) behave when being watched in two different contexts. In the first, we show that aggressive and courtship behaviors displayed by subordinate males depends critically on whether dominant males can see them, and in the second, we manipulated who was watching aggressive interactions and showed that dominant males will change their behavior depending on audience composition. In both cases, when a more dominant individual is out of view and the audience consists of more subordinate individuals, those males signal key social information to females by displaying courtship and dominant behaviors. In contrast, when a dominant male is present, males cease both aggression and courtship. These data suggest that males are keenly aware of their social environment and modulate their aggressive and courtship behaviors strategically for reproductive and social advantage.     

Mating habits of the rockfishSebastes inermis

Synopsis Underwater observations were conducted on the mating habits of rockfish,Sebastes inermis, southwest of Kyushu Island, Japan, in 1984–1986. The reproductive season of the fish lasted from October to January, when most males over 1 year of age established territories ranging from 12 to 70 m². Home ranges of females overlapped with male territories. Males performed courtship behaviors when females approached the territories. Principal motor patterns of courtship were characterized by ‘lateral display’ and ‘rushing and turning’ movements. The mating pair formed about 30 min before copulation. As the mating pair ascended to 1.5–2.0 m above the bottom, the male suddenly coiled around the female's body to copulate. In spite of frequent courtship behavior [14.8 (±10.7 SD)×15 min⁻¹, December 1985] of males, only seven copulations were observed, occurring mostly at dusk in December. In December and January, more than half of the males had accumulated urine, some large volume of it. It was assumed that the male attracted the female by releasing substances via urine during the ‘lateral display’ movements, which were performed just in front of the female, near the snout. Other functional aspects of courtship behavior are discussed.     

Do Male Veiled Chameleons,Chamaeleo calyptratus,Adjust their Courtship Displays in Response to Female Reproductive Status?

Variation in male courtship behavior may be due to inherent differences among males or may arise from males adjusting their courtship displays according to female responsiveness. Female veiled chameleons, Chamaeleo calyptratus , exhibit two distinctive suites of body coloration and behavior patterns that vary according to receptive and non-receptive stages of their reproductive cycle. We presented male chameleons with both receptive and non-receptive females, and recorded differences in their mating frequency, courtship intensity and courtship behavior patterns. As expected, males were more likely to court and attempt mating with receptive females. Although fewer males courted non-receptive females, their courtship displays were significantly longer than those directed towards receptive females. Males also adjusted the contents of their displays according to female reproductive condition. Certain behavior patterns were unique to courtship displays directed towards each class of females. Males exhibited the behavior pattern `head roll' only when paired with receptive females, and `chin rub' was displayed only during courtship of non-receptive females. We hypothesize that these differences in male courtship frequency, intensity and content reflect differences in female reproductive value. Although males may benefit from mating with both receptive and non-receptive females, the costs associated with courtship may depend on female responsiveness. Thus, males adjust their courtship tactics accordingly.     

Copulation type affects parturition in the guppy

In some animals, females are often compelled to mate with less desirable males due to the males' alternative mating tactics. Male guppies, Poecilia reticulata, exhibit courtship displays and cooperatively copulate with females. However, they also exhibit sneaking behaviors and coercively copulate with females. To examine the consequences of these two mating patterns, we investigated the influence of copulation type, i.e., cooperative or coercive, on parturition and brood size of females. A single female was allowed to freely contact and copulate with a single male only once. Males that cooperatively copulated with females had larger orange spot areas (an important criterion of female mate choice) than males that copulated coercively. Most females that were coerced into copulation did not give birth to offspring within 100 days after mating. The probability of parturition was high when females copulated cooperatively, and when their mates exhibited frequent postcopulatory jerking behavior. However, the results suggest that copulation type did not affect brood size. Brood size was positively influenced by both female body size and male orange spot area. These results suggest that parturient success is low when females are coerced into mating by less desirable males, whereas brood size is independent of copulation type.     

Cryptic forcible insemination: male snakes exploit female physiology, anatomy, and behavior to obtain coercive matings

Whether males can inseminate uncooperative females is a central determinant of mating system evolution that profoundly affects the interpretation of phenomena such as multiple mating by females, mate choice, reproductive seasonality, and courtship tactics. Forcible insemination is usually inferred from direct physical battles between the sexes and has been dismissed on intuitive grounds for many kinds of animals. For example, snakes have elongate flexible bodies (making it difficult for a male to restrain a female physically), males are typically smaller than females, and copulation requires female cloacal gaping to enable intromission. Male garter snakes (Thamnophis sirtalis) do not display any overt aggression during courtship and simply lie over the female and exhibit rhythmic pulsating caudocephalic waves of muscular contraction; previous studies have interpreted this behavior as a mechanism for eliciting female receptivity. In contrast, we show that male garter snakes forcibly inseminate females. They do so by taking advantage of specific features of snake physiology, respiratory anatomy, and antipredator behavior. The snake lung extends along most of the body, with the large posterior section (the saccular lung) lacking any respiratory exchange surface. Rhythmic caudocephalic waves by courting male garter snakes push anoxic air from the saccular lung forward and across the respiratory surfaces such that females cannot obtain oxygen. Their stress response involves cloacal gaping, which functions in other contexts to repel predators by extruding feces and musk but in this situation permits male intromission. Thus, superficially benign courtship behaviors may involve cryptic coercion even in species for which intuition dismisses any possibility of forcible insemination.     

Psychobiology of sexual behavior in a whiptail lizard, Cnemidophorus inornatus

This study investigated the effects of social environment on gonadal recrudescence and sexual behavior in male and female Little Striped Whiptail lizards (Cnemidophorus inornatus). The presence of sexually active males facilitates ovarian recrudescence in conspecific females. Similarly, the presence of reproductively active females facilitates testicular recrudescence in conspecific males. Males housed with females, however, had lower average circulating concentrations of testosterone and dihydrotestosterone, and higher average concentrations of corticosterone compared to intact males housed in isolation. In other studies, the presence of reproductively active females partially restored courtship behavior in castrated males compared to castrated males housed in isolation. Despite the stimulatory effects of females on castrates, exogenous androgens are required for complete restoration of all components of sexual behavior in male C. inornatus. Females are receptive to male courtship and copulatory behavior only during the vitellogenic stages; females in previtellogenic or postovulatory ovarian stages aggressively reject male courtship advances. These findings demonstrate reciprocal effects of sexual behaviors of males and females upon each other's reproductive behavior and physiology.     

Firefly Courtship: Behavioral and Morphological Predictors of Male Mating Success in Photinus greeni

Differences in male mating success can generate selection on male morphological traits and courtship behaviors involved in male–male competition or female mate choice. In Photinus fireflies (Coleoptera: Lampyridae), courtship is based on bioluminescent flash signals produced by both sexes. We conducted field observations of Photinus greeni fireflies engaged in competitive courtships, in which females are able to simultaneously assess several males, to identify male morphological traits and courtship behaviors that might predict male mating success. Male morphological traits did not differ between males that successfully mated compared with unsuccessful males (dialoging males that did not mate). However, courtship behavioral interactions differed: successful males tended to have higher flash pattern rates (number of flash patterns per minute), and their courtship flashes were more likely to be answered by females. We also examined how the risk of predation by Photuris fireflies altered courtship behavior of their Photinus prey. When predatory Photuris fireflies were present, P. greeni females were less likely to mate, and showed decreased flash responses to most males. However, P. greeni males that did successfully mate in spite of Photuris presence were males that maintained high flash pattern rates that elicited female responses. These results suggest that both female mate choice and Photuris predation exert strong selective pressures on the evolution of courtship signals in Photinus fireflies.     

The role of sex steroids in courtship, pairing and pairing behaviors in the socially monogamous zebra finch

The purpose of this study was to test whether sex steroid actions are necessary for courtship and pairing in socially monogamous birds. We examined the effects of an aromatase inhibitor, 1,4,6-androstatriene-3,17-dione (ATD), combined with an anti-androgen, flutamide (F), on the behavior and pairing status of initially unpaired male and female zebra finches (Taeniopygia guttata). In the first experiment, 24 adult males were implanted with either a combination of ATD and flutamide or empty implants. Two weeks after implantation, birds were housed in aviaries containing 3 ATD + F males, 3 control males, and 3 females and allowed 2 weeks to pair, with observations 7 times during the 2-week period. A second experiment tested the effects of these same treatments in females. During the first 4 days of testing, ATD + F males were less likely to attack conspecifics than were control males. ATD + F males were also less likely to "greet," or approach, females than were control males, but other courtship behaviors, including directed singing, were unaffected. ATD + F females did not differ from control females on any courtship behavior measured. Furthermore, these treatments did not affect pairing behaviors (time spent clumping or in a nest box together) or the likelihood of pairing with a partner of the opposite sex. ATD + F treatments in females did, however, increase the likelihood of same-sex pairing. This suggests that, although sex steroids may regulate some courtship behaviors in males, they do not regulate pairing behaviors and have little effect on the likelihood that a male or female will be chosen as a mate by a bird of the opposite sex.     

Visual and Olfactory Female-Borne Cues Evoke Male Courtship in the Aphid Parasitoid Aphidius colemani Viereck (Hymenoptera: Braconidae)

We investigated the courtship and mating behavior of the pan-tropical polyphagous endoparasitoid Aphidius colemani Viereck. The courtship and mating displays, the magnitude of male-male sexual approaches and the role of female-borne cues evoking male courtship behavior were quantified. The sequence of events leading to copulation in this parasitoid did not differ from that found for other braconids. Females refused to copulate more than once. Same-sex courtships were observed among males and their possible role in an adaptive context is discussed. Olfactory female-borne cues played a key role in eliciting the courtship responses of males. Males were attracted by freshly dead females, but not by dead females soaked in hexane, nor by visual cues from females alone. Intense male wing fanning behavior was elicited by crushed abdomens of virgin females, suggesting that the female abdomen is the source of a short-distance pheromone crucial in evoking male courtship. Further studies are required to clarify the exact nature of the chemicals involved.     

Testosterone response to courtship predicts future paternal behavior in the California mouse, Peromyscus californicus

In the monogamous and biparental California mouse (Peromyscus californicus), paternal care is critical for maximal offspring survival. Animals form pair bonds and do not engage in extrapair matings, and thus female evaluation of paternal quality during courtship is likely to be advantageous. We hypothesized that male endocrine or behavioral response to courtship interactions would be predictive of future paternal behavior. To test this hypothesis, we formed 20 pairs of California mice, and evaluated their behavior during the first hour of courtship interactions and again following the birth of young. We also collected blood from males at baseline, 1 hr after pairing, 3 weeks paired, and when young were 4 days old to measure testosterone (T). We found that male T-response to courtship interactions predicted future paternal behavior, specifically the amount of time he huddled over young when challenged by the temporary removal of his mate. Males that mounted T increases at courtship also approached pups more quickly during this challenge than males who had a significant decrease in T at courtship. Proximity of the male and female during courtship predicted paternal huddling during a 1-hr observation, and a multiple regression analysis revealed that courtship behavior was also predictive of birth latency. We speculate that male T-response to a female in P. californicus is an honest indicator of paternal quality, and if detectable by females could provide a basis for evaluation during mate choice.     

Mate choice in female convict cichlids (<Emphasis Type="Italic">Amatitlania nigrofasciata</Emphasis>) and the relationship between male size and dominance

We examined how male size and fighting ability influence a female’s mate assessment process and her eventual mate choice in the monogamous convict cichlid, Amatitlania nigrofasciata. Females always chose the larger of two males when they were allowed to see a larger male next to a smaller one and when a larger male defeated a smaller one in a fight. They did not differentiate between large and small males when they did not see the two males together nor did they choose a dominant over a subordinate male when both were the same size. We suggest that females select on the basis of male size because it is a better predictor of both direct and indirect benefits (i.e., future competitive interactions and foraging ability) than dominance behavior only. Despite selecting one male over the other early in the courtship period, females continued to visit both males until spawning. Our evidence suggests that this assessment behavior more closely resembles a bet-hedging tactic rather than the female’s indecision as to which male will be her mate.