Mating failure shapes the patterns of sperm precedence in an insect

Funding: University of St Andrews PhD Apprenticeship (VLB); Laidlaw Scholarship Programme (DB).Estimates of last male sperm precedence (P2) are often used to infer mechanisms of sperm competition, a form of post-copulatory sexual selection. However, high levels of mating failure (i.e. copulations resulting in no offspring) in a population can lead to misinterpretations of sperm competition mechanisms. Through simulations, García-González (2004) illustrated how mating failure could cause bimodal distributions of paternity with peaks at P2 = 0 and 1, under a random sperm mixing mechanism. Here, we demonstrate this effect empirically with the seed bug Lygaeus simulans, a species known to exhibit high levels of mating failure (40–60%), using a morphological marker to estimate paternity. Contrary to previous findings in a sister species, we did not find strong evidence for last male sperm precedence. There was a tendency towards last male precedence (P2 = 0.58) but within the expected range for random sperm mixing. Instead, P2 was highly variable, with a bimodal distribution, as predicted by García-González (2004). After taking mating failure into account, the strongest driver of paternity outcome was copulation duration. Furthermore, we found evidence that mating failure could partly be a female-associated trait. Some doubly-mated females were more likely to produce no offspring or produce offspring from two different sires than expected by chance. Therefore, some females are more prone to experience mating failure than others, a result that mirrors an earlier result in male L. simulans. Our results confirm that mating failure needs to be considered when interrogating mechanisms of post-copulatory sexual selection.Publisher PDFPeer reviewe

Can females choose to avoid mating failure in the seed bug Lygaeus simulans?

We thank the Natural Environmental Research Council (Ph.D. studentship to E.V.G.) and the Association for the Study of Animal Behaviour (Undergraduate Project Scholarship awarded to V.L.B.) for funding.It is becoming increasingly clear that copulation does not necessarily always lead to offspring production in many organisms, despite fertilization success presumably being under both strong natural and sexual selection. In the seed bug Lygaeus simulans, between 40% and 60% of copulations fail to produce offspring, with this ‘mating failure’ representing a significantly repeatable male-associated trait. Mating has been demonstrated to be costly in this species and, as such, we might expect females to minimize the chance of mating failure by displaying a preference for males with higher insemination success where possible. After assaying males for mating failure, we asked whether females preferred males with a history of successful inseminations versus unsuccessful inseminations in pairwise mate choice trials. Contrary to our expectations, females showed no preference for more successful over less successful males. Moreover, females showed no preference for larger males in the choice trials, even though larger males were significantly more likely to successfully inseminate females in the initial assay. This apparent lack of female precopulatory choice suggests that postcopulatory choice mechanisms may be key to mating failure in this species. However, this does not necessarily explain why females pay the cost of mating with males they will then reject via postcopulatory processes. More generally, our results suggest that mating failure may play a largely underappreciated role in mating systems evolution, influencing both the cost of choosiness, and the costs and benefits of polyandry.Publisher PDFPeer reviewe

The fitness effects of a pale mutant in the aposematic seed bug Lygaeus simulans indicate pleiotropy between warning coloration and life history

VLB is funded by a University of St Andrews PhD Apprenticeship. In addition, the authors would like to thank the University of St Andrews for funding open-access for this paper.Conspicuous warning colors that signal chemical or other defenses are common in the natural world. For instance, such aposematic warning patterns of red-and-black or yellow-and-black are common among insect taxa, particularly in the order Hemiptera, often forming the basis of Batesian and/or Müllerian mimicry rings. In addition, it has been repeatedly noted that color polymorphisms or mutants that influence pigmentation can show pleiotropy with other behavioral, physiological, or life-history traits. Here, we describe a pale mutant of the seed bug Lygaeus simulans that appeared in our laboratory population in 2012, which differs in color to the wild-type bugs. Through multigenerational experimental crosses between wild-type and pale mutant L. simulans, we first show that the pale phenotype segregates as a single Mendelian locus, with the pale allele being recessive to the wild type. Next, we show (a) that there is a large heterozygous advantage in terms of fecundity, (b) that pale females suffer reduced longevity, and (c) that pale males have increased body length compared to wild-type homozygotes. Our data therefore suggest that the color locus is pleiotropic with a number of life-history traits, opening the door for a more complete genetic analysis of aposematic coloration in this species. In addition, this phenotype will be useful as a visible genetic marker, providing a tool for investigating sperm competition and other post-copulatory drivers of sexual selection in this species.Publisher PDFPeer reviewe

Convalescent plasma in patients admitted to hospital with COVID-19 (RECOVERY): a randomised controlled, open-label, platform trial

SummaryBackground Azithromycin has been proposed as a treatment for COVID-19 on the basis of its immunomodulatoryactions. We aimed to evaluate the safety and efficacy of azithromycin in patients admitted to hospital with COVID-19.Methods In this randomised, controlled, open-label, adaptive platform trial (Randomised Evaluation of COVID-19Therapy [RECOVERY]), several possible treatments were compared with usual care in patients admitted to hospitalwith COVID-19 in the UK. The trial is underway at 176 hospitals in the UK. Eligible and consenting patients wererandomly allocated to either usual standard of care alone or usual standard of care plus azithromycin 500 mg once perday by mouth or intravenously for 10 days or until discharge (or allocation to one of the other RECOVERY treatmentgroups). Patients were assigned via web-based simple (unstratified) randomisation with allocation concealment andwere twice as likely to be randomly assigned to usual care than to any of the active treatment groups. Participants andlocal study staff were not masked to the allocated treatment, but all others involved in the trial were masked to theoutcome data during the trial. The primary outcome was 28-day all-cause mortality, assessed in the intention-to-treatpopulation. The trial is registered with ISRCTN, 50189673, and ClinicalTrials.gov, NCT04381936.Findings Between April 7 and Nov 27, 2020, of 16 442 patients enrolled in the RECOVERY trial, 9433 (57%) wereeligible and 7763 were included in the assessment of azithromycin. The mean age of these study participants was65·3 years (SD 15·7) and approximately a third were women (2944 [38%] of 7763). 2582 patients were randomlyallocated to receive azithromycin and 5181 patients were randomly allocated to usual care alone. Overall,561 (22%) patients allocated to azithromycin and 1162 (22%) patients allocated to usual care died within 28 days(rate ratio 0·97, 95% CI 0·87–1·07; p=0·50). No significant difference was seen in duration of hospital stay (median10 days [IQR 5 to >28] vs 11 days [5 to >28]) or the proportion of patients discharged from hospital alive within 28 days(rate ratio 1·04, 95% CI 0·98–1·10; p=0·19). Among those not on invasive mechanical ventilation at baseline, nosignificant difference was seen in the proportion meeting the composite endpoint of invasive mechanical ventilationor death (risk ratio 0·95, 95% CI 0·87–1·03; p=0·24).Interpretation In patients admitted to hospital with COVID-19, azithromycin did not improve survival or otherprespecified clinical outcomes. Azithromycin use in patients admitted to hospital with COVID-19 should be restrictedto patients in whom there is a clear antimicrobial indication

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Data from: The fitness effects of a pale mutant in the aposematic seed bug Lygaeus simulans indicate pleiotropy between warning coloration and life history

Conspicuous warning colours that signal chemical or other defences are common in the natural world. For instance, such aposematic warning patterns of red-and-black or yellow-and-black are common among insect taxa, particularly in the order Hemiptera, often forming the basis of Batesian and/or Müllerian mimicry rings. In addition, it has been repeatedly noted that colour polymorphisms or mutants that influence pigmentation can show pleiotropy with other behavioural, physiological or life-history traits. Here we describe a pale mutant of the seed bug Lygaeus simulans that appeared in our lab population in 2012, which differs in colour to the wild-type bugs. Through multigenerational experimental crosses between wild-type and pale mutant L. simulans, we first show that the pale phenotype segregates as a single Mendelian locus, with the pale allele being recessive to the wild-type. Next, we show (1) that there is a large heterozygous advantage in terms of fecundity, (2) that pale females suffer reduced longevity, and (3) that pale males have increased body length compared to wild-type homozygotes. Our data therefore suggest that the colour locus is pleiotropic with a number of life-history traits, opening the door for a more complete genetic analysis of aposematic colouration in this species. In addition, this phenotype will be useful as a visible genetic marker, providing a tool for investigating sperm competition and other post-copulatory drivers of sexual selection in this species