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Evaluation of the potential developmental toxicity of cyclododecatriene (CDDT).

The potential maternal and developmental toxicity of cyclododecatriene (CDDT) was assessed in rats. Groups of 22 time-mated female Crl:CD (SD) BR rats were exposed by inhalation (whole-body, 6 h/day) to either 0 (control), 10, 25, or 67 ppm CDDT over days 6-20 of gestation (days 6-20 G); the day of copulation plug detection was designated day 0 G. The dams were euthanized on day 21 G, and their abdominal and thoracic viscera were examined grossly. The fetuses were weighed, sexed, and examined for external, visceral, and skeletal alterations. Evidence of maternal toxicity was seen at 25 and 67 ppm. There were compound-related reductions in maternal body weight and food consumption parameters as well as increased occurrences of wet and stained fur at these exposure levels. Developmental toxicity evident as reduced mean fetal weight and delayed skeletal ossification was seen only at 67 ppm. There was no evidence of either maternal or developmental toxicity at 10 ppm. Thus, the no-observed-effect level (NOEL) for maternal toxicity was 10 ppm, and the NOEL for developmental toxicity was 25 ppm. Because developmental toxicity was observed only after exposures that also produced signs of maternal toxicity, CDDT was not considered to be a selective developmental toxicant in the rat.

Abnormalities, Drug-Induced↗

Developmental toxicity study of dimethylpiperidone.

The potential maternal and developmental toxicity of dimethylpiperidone (DMPD) was assessed in rats. Groups of 25 mated female Crl:CD (SD)IGS BR rats were exposed by inhalation (whole-body exposures) for approximately six hours per day over days 7-21 of gestation (G); day 1G was the day of copulation plug detection. The exposure levels were 0, 52, 260, or 340 (vapor plus aerosol) mg/m3 DMPD. During the in-life portion, body weights, food consumption, and clinical observation data were collected. On day 22G, the dams were euthanized and examined for gross external and internal alterations. The uterine contents were described and the fetuses were weighed and examined for external, visceral, and skeletal alterations. Maternal toxicity was seen at both 260 and 340 mg/m3. At 340 mg/m3, evidence of maternal toxicity included mortality, increased clinical observations, and decreased body weight and food consumption. At 260 mg/m3, maternal toxicity was limited to increased clinical observations and decreased food consumption. Developmental toxicity was also produced at 260 and 340 mg/m3. At 340 mg/m3, evidence of developmental toxicity included decreased fetal weight, increased embryofetal lethality with concomitant reductions in litter size, and increased fetal malformations and variations. At 260 mg/m3, effects in fetuses were limited to slightly decreased fetal weight and increased fetal variations; additionally, one litter from this level consisted entirely of resorptions. There were no compound-related effects in either dams or fetuses at 52 mg/m3. It was, therefore, concluded that DMPD was not selectively toxic to the rat conceptus.

Abnormalities, Drug-Induced↗

Beetle pollination of Philodendron solimoesense (Araceae) in French Guiana.

The pollination of Philodendron solimoesense (subgenus Meconostigma) was studied in four populations of French Guiana. Flowering is asynchronous within each population during July, and the flowering cycle is a 2-d process. Numerous insects visit Philodendron inflorescences, but the main pollinator seems to be Cyclocephala colasi (Scarabaeidae, Dynastinae). The pollination process displays aspects typical of beetle pollination: the production of heat and of a strong odor, the presence of a food reward (stigmatic secretion and sterile male flowers), and the presence of a copulation chamber. Flower heat production is important (ca. 11 degrees C above the ambient air) and may help to volatilize the fragrance. Attraction and choice-test experiments showed that C. colasi is not likely to depend on chemical information (such as pheromone) to localize conspecifics but may rely instead on stimuli produced by the inflorescences in order to meet mating partners.

Journal Article↗

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.

Animals↗

Infertile matings and sperm competition: the effect of "nonsperm representation" on intraspecific variation in sperm precedence patterns.

In theoretical and experimental approaches to the study of sperm competition, it is often assumed that ejaculates always contain enough sperm of good quality and that they are successfully transferred and used for fertilization. However, this view neglects the potential effects of infertility and sperm limitation. Permanent or temporal male infertility due to male sterility, insemination failures, or failures to fertilize the ova implies that some males do not achieve sperm representation in the female reproductive tract after mating. A review of the literature suggests that rates of nonsperm representation may be high; values for the proportion of infertile matings across 30 insect species vary between 0% and 63%, with the median being 22%. I simulated P2 (the proportion of offspring fathered by the second male to copulate with a female in a double-mating trial) distributions under a mechanism of random sperm mixing when sample sizes and rates of male infertility varied. The results show that nonsperm representation can be responsible for high intraspecific variance in sperm precedence patterns and that it can generate misleading interpretations about the mechanism of sperm competition. Nonsperm representation might be a common obstacle in the studies of sperm competition and postcopulatory female choice.

Animals↗

Male "mixed" reproductive strategies in biparental species: Trivers was probably right, but why?

Trivers proposed that, if parental care by both sexes is advantageous, males should practice a "mixed" strategy of seeking extrapair copulations, while restricting their parental investment to offspring of social mates. We explore circumstances under which males should limit their parental care in the predicted manner. We find that Trivers's "mixed" strategy will generally be evolutionarily stable so long as either socially monogamous or polygynous males usually sire more offspring per brood from a social mate than they typically sire in broods of extrapair mates. Polygynous males should spread investment across their home nests unless the expected number of chicks sired in them differs widely. Whether polygynous males should restrict paternal care to social mates' offspring hinges additionally on resident male investment in broods containing extrapair young: if resident males contribute minimally, some investment by a polygynous extrapair male becomes more advantageous. Recently reviewed data on extrapair fertilization distributions within monogamous and polygynous passerines suggest that extrapair offspring often predominate numerically within their broods, consistent with sperm expenditure theory. Nevertheless, most species conform to the model's criterion regarding relative parentage levels in broods of social versus extrapair mates. Patterns of extrapair parentage thus appear sufficient to stabilize biparental care systems.

Animals↗

The evolution of infidelity in socially monogamous passerines: neglected components of direct and indirect selection.

A recent study by Goran Arnqvist and Mark Kirkpatrick in the American Naturalist (165:S26-S37) suggested that female polyandry in birds is not driven by females because quantitative genetic approximations of selection demonstrated that indirect selection for female infidelity is weaker than natural selection against it. Instead, it was argued that extrapair copulations are the result of antagonistic selection on male behavior driving female coercion. While the approach and framework of the study were very good, the conclusions of the study were premature because a number of potential adaptive components of polyandry were unaccounted for, and several critical assumptions are unsupported by the current empirical data. Our understanding of extrapair paternity in birds, and perhaps polyandry in general, will be improved by a better empirical understanding of the direct benefits of fertility assurance and postcopulatory cryptic female choice and the relationship between polyandry and male investment. In addition, we need to develop a greater awareness of the limitations of trying to study behavior by proxy in the molecular laboratory. Together, these challenges and the framework recently presented should improve our understanding of the true function of extrapair paternity in birds.

Agonistic Behavior↗

Effects of arousal from hibernation and plasma androgen levels on mating behavior in the male big brown bat, Eptesicus fuscus.

The effects of arousal from hibernation and presence of plasma androgen on the expression of mating behavior in male big brown bats (Eptesicus fuscus) were tested in a captive population exposed to seminatural conditions in central Alabama. In the mild winter of 1994-1995, flight cage temperatures never fell below 10 degrees C. Bats were never observed to enter sustained (over 2 d) torpor. They were also never observed to mate. Unmanipulated, sham-operated, and gonadectomized males and unmanipulated females were exposed to 6 d of 4 degrees C. All individuals appeared torpid, and body temperatures of monitored bats fell at least 15 degrees-20 degrees C. Plasma androgen levels of torpid unmanipulated, sham-operated, and gonadectomized males averaged 25.4 +/- 9.2, 19.7 +/- 9.1, and 1.5 +/- 0.25 ng/mL, respectively, and did not differ significantly from levels for the same groups 1 mo previous to induced torpor. When animals were returned to 23 degrees C, 57% of unmanipulated, 40% of the sham-operated, and 33% of gonadectomized males displayed mating behavior upon arousal. Almost all matings occurred within 48 h of arousal, the majority in the first 3 h. Males not exposed to low temperatures were not observed to mate. Although individuals from all three treatments mated, gonadectomized males averaged fewer mounts and copulations per individual. Androgen levels declined significantly from torpor levels in all groups 48 h after arousal. Thus, an extended period of low body temperature and arousal appeared to be a short-term activator of sexual behavior in the big brown bat; unmanipulated males were more strongly affected by this stimulus than gonadectomized males.

Androgens↗

The pollination mechanism in Trigonidium obtusum Lindl (Orchidaceae: Maxillariinae): sexual mimicry and trap-flowers.

The pollination process in Trigonidium obtusum Lindl. (Epidendroideae: Maxillariinae) is documented. The flowers are pollinated by sexually excited drones of Plebeia droryana (Meliponinae). When attempting to copulate either with sepals or petals, these bees slip on the waxy perianth surface and become trapped in the funnel-like flower tube. Bees trying to escape from the flowers may instead access the space between the column and lip, fixing the pollinarium on their scutellum. Pollinarium-bearing bees may pollinate the flowers when repeating the above-mentioned steps, leaving pollinia on the concave stigmatic surface, thus effecting pollination. Recently removed pollinaria are too broad to enter the stigma but they begin to dehydrate and within 40 min of removal are small enough to fit the stigmatic cavity. This mechanism prevents insect-mediated self-pollination and promotes cross-pollination. Preliminary evidence based on experiments with cultivated plants suggests that they are self-compatible but that fruit set is pollinator-dependent. The data obtained are discussed in a phylogenetic context. It is suggested that the pseudocopulatory syndrome in Trigonidium could have evolved from rewardless (food advertising) ancestors. Pseudocopulation in the context of the long flowering period of this orchid species (about 7 months) is understandable since the eusocial Plebeia bees produce fertile individuals several times a year.

Animals↗

Sexual mimicry in Mormolyca ringens (Lindl.) Schltr. (Orchidaceae: Maxillariinae).

BACKGROUND AND AIMS: Pollination through sexual mimicry, also known as pseudocopulation, has been suggested to occur in some genera of the Neotropical orchid subtribe Maxillariinae. However, it has been demonstrated so far only for Trigonidium obtusum. This study reports and illustrates pollination through sexual mimicry in Mormolyca ringens. METHODS: A total of 70 h were dedicated to the observation of flowers and pollinator behaviour, which was photographically recorded. Flower features involved in pollinator attraction were studied using a stereomicroscope and by SEM analyses. Preliminary observations on the plant breeding system were made by manually self-pollinating flowers. The chemical composition of the fragrance volatiles was determined by GC/MS analysis. KEY RESULTS: The flower features of M. ringens parallel those of other pseudocopulatory flowers. The labellum shape and indument are reminiscent of an insect. Sexually excited drones of Nannotrigona testaceicornis and Scaptotrigona sp. (both in the Apidae: Meliponini) attempt copulation with the labellum and pollinate the flower in the process. In both bee species, the pollinarium is attached to the scutellum. Pollinator behaviour may promote some degree of self-pollination, but preliminary observations indicate that M. ringens flowers are self-incompatible. Flowers are produced all the year round, which ties in with the production of bee males several times a year. The phylogenetic relationships of M. ringens are discussed and a number of morphological and phenological features supporting them are reported. CONCLUSIONS: It is expected that further research could bring to light whether other Maxillariinae species are also pollinated through sexual mimicry. When a definitive and robust phylogeny of this subtribe is available, it should be possible to determine how many times pseudocopulation evolved and its possible evolutionary history.

Breeding↗

Cyclophosphamide induced DNA strand breaks in mouse embryo cephalic tissue in vivo.

Pregnant C3H mice were exposed to teratogenic doses of cyclophosphamide (CPA), on the 11th day after copulation. The effects of this agent on embryonal cephalic DNA strand breaks were assessed between 3 and 40 h after drug administration. Administration of 15, 30 and 60 mg CPA/kg body weight resulted in conversion of 23, 30 and 44% of the DNA to the single-stranded form, respectively. No detectable DNA damage was evident 3 h after drug administration, but after 6 h significant DNA damage had occurred, reaching a maximum after 9 h. However, no evidence of DNA strand breaks was present at 22, 30 and 40 h after CPA treatment, suggesting that these lesions had been repaired. These findings demonstrate that cephalic DNA damage induced by CPA in the developing mouse embryo occurs in a time and concentration dependent manner, and provide some insight into the kinetics of formation and removal of DNA strand breaks caused by CPA in vivo.

Abnormalities, Drug-Induced↗

Removal of the vomeronasal organ reduces reproductive performance and aggression in male prairie voles.

Several short-duration tests have demonstrated that the surgical removal of the vomeronasal organ (VNX) from sexually-inexperienced male rodents results in a reduction in copulatory behavior, compared to the effects of sham surgery (SHAM). We extended these studies to adult male prairie voles, Microtus ochrogaster, and substantially increased the duration of the tests. During the initial interactions with females, VNX males spent significantly less time with their noses in close proximity to the females than did SHAM males. Moreover, only two of nine VNX males sired offspring after having been paired with females for 8 weeks, whereas nine of 12 SHAM males sired offspring in that interval. We also found that VNX and SHAM males were equivalently non-aggressive to an anesthetized stimulus-male prior to being paired with females. However, after spending 2 weeks paired with a female, the VNX males were significantly less aggressive than were the SHAM males, possibly as a result of having copulated less often. In a later test, nearly all of the VNX and SHAM males that sired offspring were vigorously aggressive to a stimulus male. We conclude that the stimulation of the vomeronasal system in sexually-inexperienced male prairie voles is important for maximal reproductive performance and that the VNX-induced impairment in reproduction is associated with a decrease in inter-male aggression. The possible sensory effects of the vomeronasal system on the neural and endocrine control of reproduction and behavior are discussed.

Aggression↗

Monozygotic vs. dizygotic twin behavior in artificial mouse twins.

Adult inbred mice of an isogenic strain (AKR/NHan or C57BL/6J Han) differ in social (sexual and agonistic), emotional and psychomotoric behavior, depending on the kind of manipulation to which they were subjected at an early ontogenetic stage. Monozygotic twins (MZT) from eight-cell stages halved and transferred to uterine foster mothers were compared with dizygotic twins (DZT) from nonreduced but transferred eight-cell stages and with naturally born animals (NBA). Generally, early embryonic conditions predict the behavioral characteristics of the adult animals to a high degree. The MZT are motorially less active, less emotional, less aggressive and less socially interested than DZT and NBA. In tests of spontaneous social behavior (allogrooming, anogenital licking, mounting, fighting), as well as in tests for emotionality (open field: crossed fields and defecation), these behavioral patterns occurred less frequently in MZT than in DZT; the NBA were mostly intermediate. The copulatory pattern of male MZT differs from that of male DZT by a shortage of intromission latency and duration; furthermore, MZT pairs do not build up a steady rank order in competitive copulation tests, as opposed to DZT and NBA pairs. In a test for psychomotoric behavior (swimming), the MZT prefer "floating" as a survival strategy, whereas the DZT and NBA prefer "adult swimming." Therefore, it can be concluded that these behavioral differences may be caused by the particular psychosocial environment in which the twins grow up or may be due to early prenatal peculiarities, such as inadequate synchronization of the developmental status of uterus and embryo.

Animals↗

Reduced reproductive success for a conditioning mutant in experimental populations of Drosophila melanogaster.

Male Drosophila melanogaster that have courted newly-emerged males can modify their subsequent courtship behavior to avoid further courtship with immature males for up to 6 hr (previously reported). Here, it was hypothesized that such an experience-dependent modification would afford a mating advantage to normal males over males that carried a mutation that affects learning and memory. Coisogenic lines were constructed which varied at the dunce gene (dnc+ and dncM14 alleles) in order to test this hypothesis. Whether previously experienced with immature males or not, dnc+ and dncM14 males were indistinguishable in their response and mating efficiency when individually paired with virgin females. However, courtship performance of dnc+ and dncM14 males was different if they were first experienced with immature males and were then individually tested in an artificial population of nine immature males and one virgin female. In this situation, dnc+ males spent much less time in courtship with immature males and achieved copulation in one-third the time required for dncM14 males. As a control, the behavior and mating efficiency of courtship-naive dnc+ and dncM14 males in the artificial population was indistinguishable. In competition for a single virgin female, experienced dncM14 males showed a slight mating advantage over experienced dnc+ males. But when competition by experienced males for a single virgin female took place in the presence of nine immature males, dnc+ males were the successful maters in three-fourths of the trials.

Animals↗

Behavior and cytogenetics of fruitless in Drosophila melanogaster: different courtship defects caused by separate, closely linked lesions.

The fruitless (fru) courtship mutant was dissected into three defects of male reproductive behavior, which were separable as to their genetic etiologies by application of existing and newly induced chromosomal aberrations. fru itself is a small inversion [In(3R) 90C; 91B] on genetic and cytological criteria. Uncovering the fru distal breakpoint with deletions usually led to males with two of the fru courtship abnormalities: no copulation attempts with females (hence, behavioral sterility) and vigorous courtship among males, including the formation of "courtship chains." However, certain genetic changes involving region 91B resulted in males who formed courtship chains but who mated with females. Uncovering the fru proximal breakpoint led to males that passively elicit inappropriately high levels of courtship. This elicitation property was separable genetically from the sterility and chain formation phenotypes and provisionally mapped to the interval 89F-90F, which includes the fru proximal breakpoint. Behavioral sterility and chaining were also observed in males expressing certain abnormal genotypes, independent of the fru inversion. These included combinations of deficiencies, each with a breakpoint in 91B, and a transposon inserted in 91B.

Animals↗

Genetics of hybrid male sterility between drosophila sibling species: a complex web of epistasis is revealed in interspecific studies.

To study the genetic differences responsible for the sterility of their male hybrids, we introgressed small segments of an X chromosome from Drosophila simulans into a pure Drosophila mauritiana genetic background, then assessed the fertility of males carrying heterospecific introgressions of varying size. Although this analysis examined less than 20% of the X chromosome (roughly 5% of the euchromatic portion of the D. simulans genome), and the segments were introgressed in only one direction, a minimum of four factors that contribute to hybrid male sterility were revealed. At least two of the factors exhibited strong epistasis: males carrying either factor alone were consistently fertile, whereas males carrying both factors together were always sterile. Distinct spermatogenic phenotypes were observed for sterile introgressions of different lengths, and it appeared that an interaction between introgressed segments also influenced the stage of spermatogenic defect. Males with one category of introgression often produced large quantities of motile sperm and were observed copulating, but never inseminated females. Evidently these two species have diverged at a large number of loci which have varied effects on hybrid male fertility. By extrapolation, we estimate that there are at least 40 such loci on the X chromosome alone. Because these species exhibit little DNA-sequence divergence at arbitrarily chosen loci, it seems unlikely that the extensive functional divergence observed could be due mainly to random genetic drift. Significant epistasis between conspecific genes appears to be a common component of hybrid sterility between recently diverged species of Drosophila. The linkage relationships of interacting factors could shed light on the role played by epistatic selection in the dynamics of the allele substitutions responsible for reproductive barriers between species.

Animals↗

Bottleneck effects on genetic variance for courtship repertoire.

Bottleneck effects on evolutionary potential in mating behavior were addressed through assays of additive genetic variances and resulting phenotypic responses to drift in the courtship repertoires of six two-pair founder-flush lines and two control populations of the housefly. A simulation addressed the complication that an estimate of the genetic variance for a courtship trait (e.g., male performance vigor or the female requirement for copulation) must involve assays against the background behavior of the mating partners. The additive "environmental" effect of the mating partner's phenotype simply dilutes the net parent-offspring covariance for a trait. However, it there is an interaction with this "environmental" component, negative parent-offspring covariances can result under conditions of high incompatibility between the population's distributions for male performance and female choice requirements, despite high levels of genetic variance. All six bottlenecked lines exhibited significant differentiation from the controls in at least one measure of the parent-offspring covariance for male performance or female choice (estimated by 50 parent-son and 50 parent-daughter covariances for 10 courtship traits per line) which translated to significant phenotypic drift. However, the average effect across traits or across lines did not yield a significant net increase in genetic variance due to bottlenecks. Concerted phenotypic differentiation due to the founder-flush event provided indirect evidence of directional dominance in a subset of traits. Furthermore, indirect evidence of genotype-environment interactions (potentially producing genotype-genotype effects) was found in the negative parent-offspring covariances predicted by the male-female interaction simulation and by the association of the magnitude of phenotypic drift with the absolute value of the parent-offspring covariance. Hence, nonadditive genetic effects on mating behavior may be important in structuring genetic variance for courtship, although most of the increases in genetic variance would be expected to reflect inbreeding depression with relatively rare situations representing the facilitation of speciation by bottlenecks.

Analysis of Variance↗

Analysis of the doublesex female protein in Drosophila melanogaster: role on sexual differentiation and behavior and dependence on intersex.

doublesex (dsx) is unusual among the known sex-determination genes of Drosophila melanogaster in that functional homologs are found in distantly related species. In flies, dsx occupies a position near the bottom of the sex determination hierarchy. It is expressed in male- and female-specific forms and these proteins function as sex-specific transcription factors. In the studies reported here, we have ectopically expressed the female Dsx protein (Dsx(F)) from a constitutive promoter and examined its regulatory activities independent of other upstream factors involved in female sex determination. We show that it functions as a positive regulator of female differentiation and a negative regulator of male differentiation. As predicted by the DNA-binding properties of the Dsx protein, Dsx(F) and Dsx(M) compete with each other for the regulation of target genes. In addition to directing sex-specific differentiation, Dsx(F) plays an important role in sexual behavior. Wild-type males ectopically expressing Dsx(F) are actively courted by other males. This acquisition of feminine sex appeal is likely due to the induction of female pheromones by Dsx(F). More extreme behavioral abnormalities are observed when Dsx(F) is ectopically expressed in dsx(-) XY animals; these animals are not only courted by, but also copulate with, wild-type males. Finally, we provide evidence that intersex is required for the feminizing activities of Dsx(F) and that it is not regulated by the sex-specific splicing cascade.

Animals↗