Many species of teleost fish detect and release F prostaglandins (PGFs), but the specific identities of these compounds and how they function as species-specific pheromones have yet to be resolved. This study addressed these questions in the common carp. An initial set of experiments established that mature male common carp were attracted to chemicals released by ovulated conspecifics, whereas the odor of female goldfish, a close relative, was less attractive. Tests of fractionated holding water from ovulated carp revealed that only the non-polar fraction was attractive on its own. Mass spectrometry and immunoassay next demonstrated that the non-polar fraction contained large quantities of prostaglandin F(2α) (PGF(2α)), 15keto-prostaglandinF(2α), and 13,14-dihydro-15keto-prostaglandin F(2α) (100 g fish released over 1 μg of all 3 PGFs per h at a ratio of 1.0: 1.7: 0.7). Ovulated goldfish released the same three PGFs but at a slightly greater rate and in a different ratio. Tests of synthetic mixtures of these PGFs revealed that the carp-specific mixture attracted male carp but was no better than the goldfish-specific mixture or PGF(2α) alone and that PGF(2α) was just as attractive as mixture of all three PGFs. A final set of attraction tests revealed that although PGF(2α) could explain all of the activity of the non-polar portion of female carp holding water, it could not explain the entire activity of female water but that a mixture of PGFs and the polar fraction could. We conclude that ovulated female carp release a multi-component sex pheromone complex that is comprised of PGF(2α) and unknown species-specific polar compound(s) that synergize the activity of the former. The pheromone also might be useful in controlling this invasive species. The observation that a fish hormonal sex pheromone incorporates bodily metabolites in addition to relatively common hormonal products demonstrates a mechanism by which species specificity may be conferred to this common type of sex pheromone.
Although laboratory experiments have shown that many fishes, Goldfish (Carassius auratus) in particular, employ relatively sophisticated orientation strategies to learn the location of food in laboratory arenas, this ability has not been rigorously tested in the natural environment. In this study we documented the ability of Common Carp (Cyprinus carpio), a close relative of Goldfish, to learn the location of newly introduced food in a lake. Two experiments were conducted, the first of which determined that carp feed largely at night. The second used this information and tracked the day-and nighttime locations of 34 radio-tagged carp before and then while a food reward was introduced at a specific location in the lake for 10 days. Before the introduction of the reward, carp maintained small (∼100 m× 70 m), isolated home ranges which expanded slightly at night. This movement pattern changed after the reward was added when on the fourth night six radiotagged carp visited and exploited the reward and then returned to their home areas after sunrise. This pattern persisted for the rest of the experiment with increasing numbers of carp visiting the reward each night (21 of 34 carp visited on the tenth night) and returning to their home ranges each day. The speed and precision with which wild carp learned to exploit this reward is consistent with the social learning and spatial memory skills that they and their relatives have shown in laboratory arenas. This is particularly impressive given the turbid conditions in the lake and the lack of obvious visual landmarks.
When ovulated, female fish of many species are known to release a F-prostaglandin-derived sex pheromone that attracts conspecific males. Recently, this pheromone was identified in the common carp as a mixture of prostaglandin F(2α) (PGF(2α)) and unidentified body metabolites, which we termed a 'pheromone complex.' The present study sought to test the activity of this pheromone complex in the field by developing a system using carps implanted with PGF(2α) as pheromone donors. An initial experiment determined that osmotic pumps that delivered up to 0.4 mg of PGF(2α) per hour could be implanted into carp without any apparent effects on their health. A second experiment found that PGF(2α)-implanted male and female carp released biologically relevant (and equivalent) quantities of PGF(2α), along with two of its seemingly inactive metabolites, for up to 2 weeks. Laboratory experiments demonstrated that the odor of PGF(2α)-implanted carp was highly attractive to male conspecifics, and included necessary body metabolites; it attracted males as strongly as ovulated carp odor, and much better than PGF(2α) alone. Finally, a field test demonstrated that PGF(2α)-implanted female carp attracted mature male, but not female carp, from a distance of 20 m. This is the first demonstration of the activity of a PGF(2α)-based pheromone in a natural environment and confirms the use of a PGF-pheromone complex in the carp. We suggest that the implant technique may be useful in future studies of how PGF pheromones function and could be further developed to attract invasive fish for use in control.
We report an unusual case of communal sexual display in the arctiid moth Utetheisa ornatrix that we designate ''female pheromonal chorusing.'' As in most moths, female U. ornatrix release a long-distance sexual advertisement pheromone during a nightly activity period. We arranged U. ornatrix females in 2 types of signaling conditions: grouped and solitary. When the females were grouped with neighboring signaling females (grouped), they initiated pheromone release sooner, continued release with less interruption and over a longer total period, and performed the release with faster abdominal pumping than observed in isolated females (solitary). This differs from the usual form of sexual communication in moths: female (chemical) signalers, male receivers, and a general lack of interaction among females. At mating, male U. ornatrix transfer a large spermatophore that may enhance female reproductive success and which represents either mating effort or paternal investment. This action results in an extended postmating male refractory period leading to a female-biased operational sex ratio. We argue that this biased sex ratio generates intrasexual competition among females, to which they respond by elevating signaling effort such that the likelihood of at least matching their neighbors' signals is increased. In the field, U. ornatrix are clustered around patches of host plants, and we also explore the possibility that pheromonal chorusing is driven by cooperation among groups of related-or nonrelated-females.
Different species of water striders match leg speeds to their body sizes to maximize their jump take off velocity without breaking the water surface, which might have aided evolution of leg structures optimized for exploitation of the water surface tension. It is not understood how water striders achieve this match. Can individuals modify their leg movements based on their body mass and locomotor experience? Here we tested if water striders, Gerris latiabdominis, adjust jumping behaviour based on their personal experience and how an experimentally added body weight affects this process. Females, but not males, modified their jumping behaviour in weight-dependent manner, but only when they experienced frequent jumping. They did so within the environmental constraint set by the physics of water surface tension. Females’ ability to adjust jumping may represent their adaptation to frequent increases or decreases of the weight that they support as mating bouts, during which males ride on top of females, start or end, respectively. This suggests that natural selection for optimized biomechanics combined with sexual selection for mating adaptations shapes this ability to optimally exploit water surface tension, which might have aided adaptive radiation of Gerromorpha into a diversity of semiaquatic niches.
Perception of the female sex pheromone in Utetheisa ornatrix (Lepidoptera: Arctiidae) is responsible for induction and adjustment of calling by females and the collective phenomenon termed "female pheromonal chorusing". We found five olfactory-active compounds in the U. ornatrix female gland. When females were exposed to the entire pheromone or to two of its (synthetically prepared) components, (Z,Z,Z)-3,6,9-eicosatriene and (Z,Z,Z)-3,6,9-heneicosatriene, they were more likely to call during a given night, begin calling earlier, and briefly increase signal frequency with which they extrude their abdomen, an observable indication of calling in this species. Some females even initiated calling during photophase when exposed to the pheromone components. In general, female U. ornatrix are more sensitive to the complete blend of pheromone than to its individual compounds. We also tested the hypotheses: 1) that abdominal extrusion per se increases the rate of pheromone release; and 2) that greater abdominal pumping rhythm increases pheromone release rate. Contrary to our expectations: 1) females did not respond more strongly to a pulsed pheromone stimulus than to the constant release of pheromone at the same average release rate; and 2) we did not find a relationship between the frequency of abdominal pumping and pheromone release rate. Possible explanations for these unexpected findings are discussed.
The type II class of sex pheromones found in moths is composed of polyene hydrocarbons and their epoxides. Analysis of Utetheisa ornatrix females by gas chromatography-mass spectrometry and measurement of responses of male moths by coupled gas chromatography-electroantennogram detection confirmed the presence of large amounts of (Z,Z,Z)-1, 3,6,3,6,[9][10][11][12][13][14][15][16][17][18][19][20][21] and smaller amounts of (Z,Z,Z)-3,6,9-heneicosatriene (3,6,9-21:Hy). Both compounds were detected in pheromone glands of newly emerged adults, with low amounts found in the late pupal stage, indicating that sex pheromone biosynthesis started in the late pupal stage. In our population of females (several hundred sampled), approximately 90% produced the tetraene, 1,3,6,9-21:Hy, as the major component, while the other 10% produced only a large amount (1500-2000 ng) of 3,6,9-21:Hy, with no detectable amount of the tetraene. This result could indicate that two distinct populations are present in our original collection site in Florida. Decapitated female moths accumulated 3,6,9-21:Hy and 1,3,6,9-21:Hy compared to the same age normal females, indicating that female moths continuously produce pheromone. A pheromone biosynthesis activating neuropeptide (PBAN)-like neuropeptide did not affect sex pheromone production as indicated by injection of synthetic PBAN and decapitation of U. ornatrix female adults. When the labeled precursor, D4-9,12,15-18:acid, was injected into the early pupal stage, the most abundantly labeled hydrocarbons were 3,6,9-21:Hy and 1,3,6,9-21:Hy in the female adults. This result indicated that 3,6,9-21:Hy could be biosynthesized from linolenic acid through chain elongation and decarboxylation. To determine how 1,3,6,9-21:Hy is produced, D4-3,6,9-21:Hy was injected into pupae and monitored for incorporation of label. No label was incorporated into 1,3,6,9-21:Hy, although a large amount of triene, 3,6,9-21:Hy, was recovered in the pheromone gland. This indicates that U. ornatrix females do not use 3,6,9-21:Hy to produce 1,3,6,9-21:Hy, and the terminal double bond is introduced earlier in the biosynthetic pathway. RightsWorks produced by employees of the U.S. Government as part of their official duties are not copyrighted within the U.S. The content of this document is not copyrighted. Abstract The type II class of sex pheromones found in moths is composed of polyene hydrocarbons and their epoxides. Analysis of Utetheisa ornatrix females by gas chromatography-mass spectrometry and measurement of responses of male moths by coupled gas chromatography-electroantennogram detection confirmed the presence of large amounts of (Z,Z,Z)-1,3,6,9-heneicosatetraene (1,3,6,9-21:Hy) and smaller amounts of (Z,Z, Z)-3,6,9-heneicosatriene (3,6,9-21:Hy). Both compounds were detected in pheromone glands of newly emerged adults, with low amounts found in the late pupal stage, indicating that sex pheromone biosynthesis started in the late pupal stage. In our population of females (several hundred sampled), approximately ...
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