A single synonymous nucleotide change impacts the male-killing phenotype of prophage WO gene wmk

Wolbachia are the most widespread bacterial endosymbionts in animals. Within arthropods, these maternally-transmitted bacteria can selfishly hijack host reproductive processes to increase the relative fitness of their transmitting females. One such form of reproductive parasitism called male killing, or the selective killing of infected males, is recapitulated to degrees by transgenic expression of the WO-mediated killing (wmk) gene. Here, we characterize the genotype-phenotype landscape of wmk-induced male killing in D. melanogaster using transgenic expression. While phylogenetically distant wmk homologs induce no sex-ratio bias, closely-related homologs exhibit complex phenotypes spanning no death, male death, or death of all hosts. We demonstrate that alternative start codons, synonymous codons, and notably a single synonymous nucleotide in wmk can ablate killing. These findings reveal previously unrecognized features of transgenic wmk-induced killing and establish new hypotheses for the impacts of post-transcriptional processes in male killing variation. We conclude that synonymous sequence changes are not necessarily silent in nested endosymbiotic interactions with life-or-death consequences.

A single synonymous nucleotide change impacts the male-killing phenotype of prophage WO gene wmk

Wolbachia are the most widespread bacterial endosymbionts in animals. Within arthropods, these maternally-transmitted bacteria can selfishly hijack host reproductive processes to increase the relative fitness of their transmitting females. One such form of reproductive parasitism called male killing, or the selective killing of infected males, is recapitulated to degrees by transgenic expression of the WO-mediated killing gene wmk. Here, we characterize the genotype-phenotype landscape of wmk-induced male killing in D. melanogaster. While phylogenetically distant wmk homologs induce no sex-ratio bias, closely-related homologs exhibit complex phenotypes spanning no death, male death, or death of all hosts. We demonstrate that alternative start codons and, notably, one synonymous mutation in wmk can ablate killing. These findings reveal previously unrecognized relationships of wmk-induced killing and establish new hypotheses for the impacts of post-transcriptional processes in wmk-induced male killing. We conclude that single synonymous sequence changes are not necessarily silent in important nested symbiotic interactions.

X-linked meiotic drive can boost population size and persistence

X-linked meiotic drivers cause X-bearing sperm to be produced in excess by male carriers, leading to female-biased sex ratios. Here, we find general conditions for the spread and fixation of X-linked alleles. Our conditions show that the spread of X-linked alleles depends on sex-specific selection and transmission rather than the time spent in each sex. Applying this logic to meiotic drive, we show that polymorphism is heavily dependent on sperm competition induced both by female and male mating behavior and the degree of compensation to gamete loss in the ejaculate size of drive males. We extend these evolutionary models to investigate the demographic consequences of biased sex ratios. Our results suggest driving X-alleles that invade and reach polymorphism (or fix and do not bias segregation excessively) will boost population size and persistence time by increasing population productivity, demonstrating the potential for selfish genetic elements to move sex ratios closer to the population-level optimum. However, when the spread of drive causes strong sex-ratio bias, it can lead to populations with so few males that females remain unmated, cannot produce offspring, and go extinct. This outcome is exacerbated when the male mating rate is low. We suggest that researchers should consider the potential for ecologically beneficial side effects of selfish genetic elements, especially in light of proposals to use meiotic drive for biological control.

Spatial and temporal sex ratio bias and Wolbachia-infection in New Zealand Crambidae (Lepidoptera: Pyraloidea)

The New Zealand fauna of snout moths (Pyraloidea) predominantly consists of endemic species. During 2017 and 2018, 56 species of Pyraloidea in 1,749 individuals were collected at 14 localities. All species were screened for Wolbachia-infection, with specimens of eight species (14%) being positive, of which six species belong to Scopariinae. This is the first record of Wolbachia-infection amongst New Zealand Lepidoptera. The most common pyraloid species, Eudonia submarginalis and Orocrambus flexuosellus, were analysed for a larger set of individuals looking for sex ratio and Wolbachia-infection. There is a sex ratio bias towards females in both species, but it varies in space and time. Wolbachia is found in all populations of E. submarginalis with 10–80% of the tested individuals being positive, depending on locality. No Wolbachia-infection has been found in O. flexuosellus. Thus, sex ratio bias might be linked to Wolbachia-infection in E. submarginalis, but not in O. flexuosellus.

Sexual dimorphism and sex ratio bias in the dioecious willow Salix purpurea L

PremiseSexual dimorphism in dioecious plant species is often not obvious or is absent. Dioecious species populations also often exhibit deviations from expected sex ratios. Previous studies on members of the Salicaceae family have shown strong, partial, and no sexual dimorphism. Some studies have shown sex-biased ratios in several Salix spp., however, S. purpurea has never been examined for evidence of sexual dimorphism or for the presence of sex-ratio bias, and therefore a comprehensive phenotypic study is needed to fill this knowledge gap.MethodsThis study examined a suite of morphological, phenological, physiological and wood composition traits from multi-environment and multi-year replicated field trials in a diversity panel of unrelated S. purpurea accessions and in full-sib F1 and F2 families produced through controlled cross pollinations to test for sexual dimorphism and sex ratio bias.Key ResultsSignificant evidence of sexual dimorphism was found in vegetative traits with greater means for many traits in male genotypes compared to females across three populations of S. purpurea, measured across multiple years that were highly predictive of biomass yield. Male plants exhibited greater nitrogen accumulation under fertilizer amendment as measured by SPAD in the diversity panel, and males showed greater susceptibility to fungal infection by Melampsora spp in the F2 family. There were also consistent female-biased sex ratios in both the F1 and F2 families.ConclusionsThese results provide the first evidence of sexual dimorphism in S. purpurea and also confirm the prevalence of female-biased sex ratios previously found in other Salix species.

Persistence of a Wolbachia-driven sex ratio bias in an island population of Eurema butterflies

It is generally believed that when maternally inherited sex ratio distorters become predominant, either the host population goes extinct or nuclear suppressors evolve in the host. Here, we show an empirical case where all-female-producing Wolbachia is likely to be stably maintained at a high frequency. On an island population of the butterfly Eurema mandarina, a Wolbachia strain wFem, which makes female hosts produce all-female offspring without sibling lethality (female drive), is highly prevalent. We found that, with some fluctuations, wFem appeared to be stably maintained for at least 12 years at a high frequency, resulting in the existence of an abnormally high number of virgin females. Interestingly, comparison between sex ratios of captive individuals and sex ratios deduced from wFem frequencies suggested a plastic behavioral change of males and females in response to the shift of sex ratios. wFem presence does not affect brood size but has a slightly negative effect on body size. Stable coexistence of wFem-positive and -negative females in the population may be explained via mate choice by males, which keeps wFem in check. Taken together, this butterfly population is an attractive model for future studies on the population dynamics of sex ratios and mating behavior.

Social structure and demography of a remnant Asian elephant Elephas maximus population and the implications for survival

The Asian elephant Elephas maximus is at risk of extinction as a result of anthropogenic pressures, and remaining populations are often small and fragmented remnants, occupying a fraction of the species' former range. Once widely distributed across China, only a maximum of 245 elephants are estimated to survive across seven small populations. We assessed the Asian elephant population in Nangunhe National Nature Reserve in Lincang Prefecture, China, using camera traps during May–July 2017, to estimate the population size and structure of this genetically important population. Although detection probability was low (0.31), we estimated a total population size of c. 20 individuals, and an effective density of 0.39 elephants per km2. Social structure indicated a strong sex ratio bias towards females, with only one adult male detected within the population. Most of the elephants associated as one herd but three adult females remained separate from the herd throughout the trapping period. These results highlight the fragility of remnant elephant populations such as Nangunhe and we suggest options such as a managed metapopulation approach for their continued survival in China and more widely.

Female bias in an immigratory population of Cnaphalocrocis medinalis moths based on field surveys and laboratory tests

Sex ratio bias is common in migratory animals and can affect population structure and reproductive strategies, thereby altering population development. However, little is known about the underlying mechanisms that lead to sex ratio bias in migratory insect populations. In this study, we used Cnaphalocrocis medinalis, a typical migratory pest of rice, to explore this phenomenon. A total of 1,170 moths were collected from searchlight traps during immigration periods in 2015–2018. Females were much more abundant than males each year (total females: total males = 722:448). Sex-based differences in emergence time, take-off behaviour, flight capability and energy reserves were evaluated in a laboratory population. Females emerged 0.78 days earlier than males. In addition, the emigratory propensity and flight capability of female moths were greater than those of male moths, and female moths had more energy reserves than did male moths. These results indicate that female moths migrate earlier and can fly farther than male moths, resulting more female moths in the studied immigratory population.