Sex in the wild: How and why field‐based studies contribute to solving the problem of sex*

Why and how sexual reproduction is maintained in natural populations, the so‐called “queen of problems,” is a key unanswered question in evolutionary biology. Recent efforts to solve the problem of sex have often emphasized results generated from laboratory settings. Here, we use a survey of representative “sex in the wild” literature to review and synthesize the outcomes of empirical studies focused on natural populations. Especially notable results included relatively strong support for mechanisms involving niche differentiation and a near absence of attention to adaptive evolution. Support for a major role of parasites is largely confined to a single study system, and only three systems contribute most of the support for mutation accumulation hypotheses. This evidence for taxon specificity suggests that outcomes of particular studies should not be more broadly extrapolated without extreme caution. We conclude by suggesting steps forward, highlighting tests of niche differentiation mechanisms in both laboratory and nature, and empirical evaluation of adaptive evolution‐focused hypotheses in the wild. We also emphasize the value of leveraging the growing body of genomic resources for nonmodel taxa to address whether the clearance of harmful mutations and spread of beneficial variants in natural populations proceeds as expected under various hypotheses for sex.     

High‐performance liquid chromatography quantification of enantiomers of a Dihydroxylated tetrahydrofuran natural product

Both enantiomers of petromyroxol are putative pheromones in sea lamprey (Petromyzon marinus). Here, we describe the separation and quantification of the petromyroxol enantiomers using high‐performance liquid chromatography tandem mass spectrometry. The separation was tested on a wide range of chiral columns with normal phases, and effects of the chromatographic parameters such as mobile phase and temperature on the separation were optimized. The AD‐H column showed the best separation of enantiomers with n‐hexane and ethanol as the mobile phase. The enantiomers were detected by multiple reaction monitoring with a positive atmospheric‐pressure chemical ionization on triple quadrupole mass spectrometer. Validation revealed that the method was specific, accurate, and precise. The validated method was applied to measure the amount of petromyroxol enantiomers in water conditioned with sea lamprey larvae, the source of the putative pheromone. This method will be applied in quantifying the natural scalemic petromyroxol mixture, enabling further investigations of a rare non‐racemic enantiomeric pheromone mixture in a vertebrate species.     

The courtship behaviour of Yponomeuta padellus

The sequence of behavioural steps from male attraction until copulation is described and analyzed for Yponomeuta padellus L. (Yponomeutidae, Lepidoptera). Males respond to a pheromone emitting female with wing fanning followed by upwind zigzagging flight. Males will land near the females and then walk while wing fanning. The antennae drum onto the substratum and the abdominal-hair pencils are displayed. When in contact with a female the male drums with his antennae onto wings, head and antennae. Mostly a clasping attempt follows. A male has a higher chance to be accepted when he firt encounters the rear of a female instead of the head. Females refuse males by walking away or wing fanning or a combination of both.

---

Résumé Les séquences du comportement depuis l'attraction du mâle jusqu'à la copulation ont été décrites et analysées chez Y. padellus (L.). Les mâles réagissent à une femelle émenttant une phéromone par un hattement d'ailes suivi d'un vol en zigzag contre le vent. Les mâles atterriront près des femelles et marcheront avec leurs ailes en battement. Les antennes tambourinent le substrat et les pinceaux abdominaux sont déplyés. Lorsqu'il est au contact de la femelle, le mâle tambourine avec ses antennes les ailes, la tête et les antennes. Généralement suit une tentative d'étreinte. Les chances de succès sont les plus élevées quand les premiers contacts du mâle sont avec la partie postérieure de la femelle plutôt qu'avec la tête. Les femelles refusent les mâles en s'écartant ou en battant des ailes ou les deux à la fois.Il semble qu'en plus des stimuli cliniques, des stimuli mécaniques fournis par les écailles sont nécessaires pour que les mâles localisent la femelle et s'orientent pour une tentative d'étreinte.

     

Identification of aggregation substances of Enterococcus faecalis cells after induction by sex pheromones

The sex pheromone system of Enterococcus faecalis is responsible for the clumping response of a plasmid carrying donor strain with a corresponding plasmid free recipient strain due to the production of sex pheromones by the recipient strain. The clumping response is mediated by a surface material (called aggregation substance) which is synthesized upon addition of sex pheromones to the cultures. Here we show that after induction a dense layer of hairlike structures is formed on the cell wall of the bacteria. These hairlike structures are responsible for the cell-cell contact which leads to the aggregation of cells. Formation of these structures was specific, only occurring after the addition of homologous sex pheromone.Abbreviations cAD1 sex pheromone specific for plasmid pAD1 - cPD1 sex pheromone specific for plasmid pPD1 - CW cell wall - pAD1 conjugative plasmid specifically transferred in the presence of cAD1 - pPD1 conjugative plasmid specifically transferred in the presence of cPD1 - PBS 10 mM Na-phosphate pH 7.5, 0.85% NaCl     

An analysis of anemotactic flight in female moths stimulated by host odour and comparison with the males' response to sex pheromone

ABSTRACT. The flight pattern of mated female navel orangeworm moths, Amyelois transitella (Walker), responding to odour from potential larval hosts is zigzagging upwind flight. However, at times these moths are capable of flying nearly directly upwind towards the odour source (track angles near 0). This response indicates that these females are capable of very accurate anemotactic control of their heading or course angle, since small angular errors in this measure would translate into larger deviations from direct upwind flight. Males of this species exhibit flight patterns similar to those of females, including track angles clustered about 0 when flying upwind to a source of the female-produced pheromone, but under these experimental conditions they flew with a higher average airspeed than the females. When females lose contact with an odour plume they initiate a well-defined programme of cross-wind counterturning or casting, which may normally increase their chances of retrieving contact with that plume when the wind direction shifts. The resultant track angles of females increase significantly by 0.8 s after plume loss, indicating that the female has initiated changes in both her course angle and airspeed. By 1 s after plume loss the females' track angles are no longer unimodally distributed about 0, but are bimodally distributed about -90 and +90. Males responded more rapidly to the loss of a pheromone plume, demonstrating a significant change in track angle 0.4 s after plume loss. Overall, female and male A.transitella exhibited remarkably similar anemotactic flight manoeuvres during upwind flight to odour sources as well as after plume loss.     

Endocrine regulation of female contact sex pheromone production in the German cockroach, Blattella germanica

ABSTRACT The amount of the major component of the cuticular sex pheromone, 3, 11-dimethyl-2-nonacosanone, on individual female German cockroaches, Blattella germanica (L.), as a function of age was determined by gas-liquid-chromatographic analysis. Accumulation of phermone increased with age in both virgin and mated females. During the first gono-trophic cycle, the pheromone accumulated most rapidly when oocyte growth rates were maximal (days 5–10), and least rapidly while the female carried an ootheca (days 11–32). Pheromone accumulation was similar in virgin and mated females when the same physiological stages (determined by oocyte maturation) were considered. Inhibition of Juvenile Hormone release, through allatectomy, chemicals (precocene or fluoromevalonate), or through mechanical egg case implants, suppressed or delayed pheromone production and oocyte growth. The Juvenile Hormone analogue ZR512 induced allatectomized or head-ligated females and females with chemically or mechanically inhibited corpora allata to produce pheromone and enlarge their basal oocytes. Finally, ZR512 applied to intact females stimulated pheromone production in a dose-dependent manner.     

Selection for altered pheromone-component ratios in the pink bollworm moth,Pectinophora gossypiella (Lepidoptera: Gelechiidae)

Female pink bollworm moths, Pectinophora gossypiella(Saunders), were selected for altered component ratios of the long-distance sex pheromone, (Z, E)-and (Z, Z)-7,11- hexadecadienyl acetate. Selection for 12 generations increased the mean (± SD) percentage of the (Z, E)isomer from 42.9 ± 1.0 in the parental generation to 48.2 ± 1.2. Although statistically significant, a change of this magnitude may be transparent to males because of their relatively broad response spectrum. Selection for a lower percentage of the (Z, E)isomer yielded no change in the mean pheromone ratio. The total amount of pheromone produced declined in both selected lines. In the line selected for females producing a high percentage of the (Z, E)isomer, the duration of wing fanning by males to high (Z, E)blends was elevated.     

Competition for hosts between two hyperparasitoids of aphids,Dendrocerus laticeps andDendrocerus carpenteri (Hymenoptera: Megaspilidae): The benefit of interspecific host discrimination

The two hyperparasitoids of aphids, Dendrocerus carpenteri(Curtis) and Dendrocerus laticeps(Hedicke), differ in their interspecific host discrimination behavior: D. carpenterireadily accepts hosts for oviposition previously parasitized by D. laticeps,whereas D. laticepsoften refrains from ovipositing on hosts previously parasitized by D. carpenteri.The two species also differ in their competitive abilities: D. carpenteriis competitively superior to D. laticepswhen the time between ovipositions is 1 h. D. carpenteri's superiority probably arises from (a) the action of the venom produced by the female to paralyze the host, as evidenced by the comparison of emergence patterns from naturally vs artificially parasitized hosts, and (b) its more rapid egg development. Because of its competitive abilities, D. carpenterigains only limited benefit from discriminating interspecifically, at least when encountering a host parasitized by a competitively inferior species, such as D. laticeps.In contrast, D. laticepsgains considerable benefit from recognizing hosts previously parasitized by D. carpenteri,as the probability to produce offspring from such hosts is greatly reduced.     

Disruption of pheromone communication in moths: is the natural blend really most efficacious?

It is widely assumed that a blend of pheromone components, that is qualitatively and quantitatively similar to the natural sex attractant, is the most effective mating disruptant for moths. However, the literature contains only limited evidence supporting this assumption. The authors discuss the importance of comparing the relative effectiveness of complete and incomplete pheromone blends when evaluating the potential of mating disruption for controlling pest moths.     

Biomechanics of the release of sex pheromone in moths: effects of body posture on local airflow

ABSTRACT. The calling posture of Spilosoma congrua Walker (Lepidoptera: Arctiidae) affects local airflow in ways that may impact on the release of chemical signals. Dried specimens of S. congrua imitating resting moths, calling moths, and moths with intermediate postures were placed in a wind tunnel. The wind speed was measured near the abdomenal tip where sex pheromone emission occurs. The calling posture, wings-up, gave significantly greater mean wind speeds near the surface of the sex pheromone gland, effectively eliminating the dead air space that surrounds a non-calling moth. The calling posture also decreased the relative intensity of turbulence near the pheromone gland surface.