Selection on female reproductive schedules in the marula fly, Ceratitis cosyra (Diptera: Tephritidae) affects dietary optima for female reproductive traits but not lifespan.

Introduction: A changing environment can select on life-history traits and trade-offs in a myriad of ways. For example, global warming may shift phenology and thus the availability of host-plants. This may alter selection on survival and fertility schedules in herbivorous insects. If selection on life-histories changes, this may in turn select for altered nutrient intake, because the blend of nutrients organisms consume helps determine the expression of life-history traits. However, we lack empirical work testing whether shifts in the timing of oviposition alter nutrient intake and life-history strategies. Methods: We tested in the marula fruit fly, Ceratitis cosyra, how upward-selection on the age of female oviposition, in comparison with laboratory adapted control flies, affects the sex-specific relationship between protein and carbohydrate intake and life-history traits including lifespan, female lifetime egg production and daily egg production. We then determined the macronutrient ratio consumed when flies from each selection line and sex were allowed to self-regulate their intake. Results: Lifespan, lifetime egg production and daily egg production were optimised at similar protein to carbohydrate (P:C) ratios in flies from both selection lines. Likewise, females and males of both lines actively defended similar nutrient intake ratios (control =1:3.6 P:C; upward-selected = 1:3.2 P:C). Discussion: Our results are comparable to those in non-selected C. cosyra, where the optima for each trait and the self-selected protein to carbohydrate ratio observed were nearly identical. The nutrient blend that needs to be ingested for optimal expression of a given trait appeared to be well conserved across laboratory adapted and experimentally selected populations. These results suggest that in C. cosyra, nutritional requirements do not respond to a temporal change in oviposition substrate availability.

Ovarian apoptosis is regulated by carbohydrate intake but not by protein intake in speckled cockroaches.

When the likelihood of reproducing successfully is low, any prior investment in developing oocytes may be wasted. One means of recouping this investment is oosorption - where ova are absorbed and resources salvaged so they can be re-allocated to other traits. Food-limited female speckled cockroaches (Nauphoeta cinerea) appear to use this strategy. However, it is unclear if total food intake or the availability of specific nutrients induces this process. Here, we used the geometric framework of nutrition to determine how protein, carbohydrate and energy intake affect levels of ovarian apoptosis and necrosis (controlled versus uncontrolled cell death) in the terminal oocytes of female N. cinerea. We then compare the effects of nutrient intake on apoptosis (a key step towards oosorption) and offspring production to better understand the relationship between diet, apoptosis and female fitness. We found that even when food was abundant, females experienced high levels of apoptosis if their diet lacked carbohydrate. Necrosis was reduced when energy intake was high, but largely irrespective of nutrient ratio. Offspring production peaked on a low protein, high carbohydrate nutrient ratio (1P:7.96C), similar to that which minimized apoptosis (1P:7.34C) but not in the region of nutrient space that minimized necrosis. Thus, females consuming an ideal nutrient blend for reproduction can invest heavily in their current brood without needing to salvage nutrients from developing ova. However, offspring production was more dependent on carbohydrate consumption than apoptosis was, suggesting that the importance of carbohydrate in reproduction goes beyond regulating oosorption. This reliance on carbohydrate for female reproduction may reflect the unusual reproductive and nutritional physiology of speckled cockroaches; attributes that make this species an exciting model for understanding how diet regulates reproduction.

A sex skew in life-history research: the problem of missing males.

Life-history strategies are diverse. While understanding this diversity is a fundamental aim of evolutionary biology and biodemography, life-history data for some traits-in particular, age-dependent reproductive investment-are biased towards females. While other authors have highlighted this sex skew, the general scale of this bias has not been quantified and its impact on our understanding of evolutionary ecology has not been discussed. This review summarizes why the sexes can evolve different life-history strategies. The scale of the sex skew is then discussed and its magnitude compared between taxonomic groups, laboratory and field studies, and through time. We discuss the consequences of this sex skew for evolutionary and ecological research. In particular, this sex bias means that we cannot test some core evolutionary theory. Additionally, this skew could obscure or drive trends in data and hinder our ability to develop effective conservation strategies. We finally highlight some ways through which this skew could be addressed to help us better understand broad patterns in life-history strategies.

Oxidative Damage Is Influenced by Diet But Unaffected by Selection for Early Age of Oviposition in the Marula Fly, Ceratitis cosyra (Diptera: Tephritidae).

The expression of life-history traits, such as lifespan or reproductive effort, is tightly correlated with the amount and blend of macronutrients that individuals consume. In a range of herbivorous insects, consuming high protein to carbohydrate ratios (P:C) decreases lifespan but increases female fecundity. In other words, females face a resource-based trade-off between lifespan and fecundity. Redox metabolism may help mediate this trade-off, if oxidative damage is elevated by reproductive investment and if this damage, in turn, reduces lifespan. Here, we test how diets varying in P:C ratio affect oxidative damage and antioxidant protection in female and male of the marula fly, Ceratitis cosyra (Diptera: Tephritidae). We use replicated lines that have been subjected to experimental evolution and differ in their lifespan and reproductive scheduling. We predicted that high fecundity would be associated with high oxidative damage and reduced antioxidant defences, while longer lived flies would show reduced damage and elevated antioxidant defences. However, higher levels of oxidative damage were observed in long-lived control lines than selection lines, but only when fed the diet promoting lifespan. Flies fed diets promoting female fecundity (1:4 and 1:2 P:C) suffered greater oxidative damage to lipids than flies fed the best diet (0:1 P:C) for lifespan. Total antioxidant capacity was not affected by the selection regime or nutrition. Our results reiterate the importance of nutrition in affecting life-history traits, but suggest that in C. cosyra, reactive oxygen species play a minimal role in mediating dietary trade-offs between lifespan and reproduction.

Mapping sex differences in the effects of protein and carbohydrates on lifespan and reproduction in Drosophila melanogaster: is measuring nutrient intake essential?

Understanding how diet affects reproduction and survival is a central aim in evolutionary biology. Although this relationship is likely to differ between the sexes, we lack data relating diet to male reproductive traits. One exception to this general pattern is Drosophila melanogaster, where male dietary intake was quantified using the CApillary FEeder (CAFE) method. However, CAFE feeding reduces D. melanogaster survival and reproduction, so may distort diet-fitness outcomes. Here, we use the Geometric Framework of Nutrition to create nutrient landscapes that map sex-specific relationships between protein, carbohydrate, lifespan and reproduction in D. melanogaster. Rather than creating landscapes with consumption data, we map traits onto the nutrient composition of forty agar-based diets, generating broad coverage of nutrient space. We find that male and female lifespan was maximised on low protein, high carbohydrate blends (~ 1P:15.9C). This nutrient ratio also maximised male reproductive rates, but females required more protein to maximise daily fecundity (1P:1.22C). These results are consistent with CAFE assay outcomes. However, the approach employed here improved female fitness relative to CAFE assays, while effects of agar versus CAFE feeding on male fitness traits depended on the nutrient composition of experimental diets. We suggest that informative nutrient landscapes can be made without measuring individual nutrient intake and that in many cases, this may be preferable to using the CAFE approach. The most appropriate method will depend on the question and species being studied, but the approach adopted here has the advantage of creating nutritional landscapes when dietary intake is hard to quantify.

Complex relationship between amino acids, fitness and food intake in Bombus terrestris.

The ratio of amino acids to carbohydrates (AA:C) that bumble bees consume has been reported to affect their survival. However, it is unknown how dietary AA:C ratio affects other bumble bee fitness traits (e.g., fecundity, condition) and possible trade-offs between them. Moreover, while individual AAs affect phenotype in many species, the effects of AA blend on bumble bee fitness and food intake are unclear. We test how the AA:C ratio that bumble bees (Bombus terrestris) consume affects their condition (abdomen lipid and dry mass), survival following food removal, and ovarian activation. We then compare ovarian activation and food intake in bees fed identical AA:C ratios, but where the blend of AAs in diets differ, i.e., diets contained the same 10 AAs in an equimolar ratio or in the same ratio as in bee collected pollen. We found that AA:C ratio did not significantly affect survival following food removal or ovarian activation; however, high AA intake increased body mass, which is positively correlated with multiple fitness traits in bumble bees. AA blend (i.e., equimolar versus pollen) did not significantly affect overall ovarian activation or consumption of each experimental diet. However, there was an interaction between AA mix and dietary AA:C ratio affecting survival during the feeding experiment, and signs that there may have been weak, interactive effects of AA mix and AA:C ratio on food consumption. These results suggest that the effect of total AA intake on bumble bee phenotype may depend on the blend of individual AAs in experimental diets. We suggest that research exploring how AA blend affects bumble bee performance and dietary intake is warranted, and highlight that comparing research on bee nutrition is complicated by even subtle variation in experimental diet composition.

The myriad of complex demographic responses of terrestrial mammals to climate change and gaps of knowledge: A global analysis.

Approximately 25% of mammals are currently threatened with extinction, a risk that is amplified under climate change. Species persistence under climate change is determined by the combined effects of climatic factors on multiple demographic rates (survival, development and reproduction), and hence, population dynamics. Thus, to quantify which species and regions on Earth are most vulnerable to climate-driven extinction, a global understanding of how different demographic rates respond to climate is urgently needed. Here, we perform a systematic review of literature on demographic responses to climate, focusing on terrestrial mammals, for which extensive demographic data are available. To assess the full spectrum of responses, we synthesize information from studies that quantitatively link climate to multiple demographic rates. We find only 106 such studies, corresponding to 87 mammal species. These 87 species constitute <1% of all terrestrial mammals. Our synthesis reveals a strong mismatch between the locations of demographic studies and the regions and taxa currently recognized as most vulnerable to climate change. Surprisingly, for most mammals and regions sensitive to climate change, holistic demographic responses to climate remain unknown. At the same time, we reveal that filling this knowledge gap is critical as the effects of climate change will operate via complex demographic mechanisms: a vast majority of mammal populations display projected increases in some demographic rates but declines in others, often depending on the specific environmental context, complicating simple projections of population fates. Assessments of population viability under climate change are in critical need to gather data that account for multiple demographic responses, and coordinated actions to assess demography holistically should be prioritized for mammals and other taxa.

Herbaceous perennial plants with short generation time have stronger responses to climate anomalies than those with longer generation time.

There is an urgent need to synthesize the state of our knowledge on plant responses to climate. The availability of open-access data provide opportunities to examine quantitative generalizations regarding which biomes and species are most responsive to climate drivers. Here, we synthesize time series of structured population models from 162 populations of 62 plants, mostly herbaceous species from temperate biomes, to link plant population growth rates (λ) to precipitation and temperature drivers. We expect: (1) more pronounced demographic responses to precipitation than temperature, especially in arid biomes; and (2) a higher climate sensitivity in short-lived rather than long-lived species. We find that precipitation anomalies have a nearly three-fold larger effect on λ than temperature. Species with shorter generation time have much stronger absolute responses to climate anomalies. We conclude that key species-level traits can predict plant population responses to climate, and discuss the relevance of this generalization for conservation planning.

Dose-dependent effects of antibiotic intake on Bombus Terrestris (Linnaeus, 1758) dietary intake, survival and parasite infection prevalence.

Diseases may contribute to the widespread declines seen in many bee species. The gut bacteria of bees may serve as one defence against disease, by preventing pathogen colonisation. However, exposure to antibiotics on forage or in the hive may disrupt bee gut bacteria and remove this protective effect. A number of studies show that high antibiotic doses reduce bee health but the effects of field-realistic antibiotic doses remain unclear. Here, we test how Bombus terrestris (Linnaeus, 1758) is affected by multiple field-realistic concentrations of the antibiotic oxytetracycline, which is sometimes used to protect flowering crops from bacterial infections. We measured survival, feeding behaviour and the likelihood of developing infection with the gut parasitic trypanosome Crithidia bombi Lipa & Triggiani, 1988 following oral inoculation with a range of antibiotic doses. Rising antibiotic concentrations were associated with reduced survival and food consumption, and an increased likelihood of becoming infected with C. bombi. These effects were seen at antibiotic concentrations that are applied to crops and so may be encountered by foraging bees in the field. These results support the hypothesis that field-realistic antibiotic doses have lethal and sub-lethal effects on B. terrestris and highlight the importance of improving our understanding of how field-realistic antibiotic doses affect pollinators.

Offspring sex ratios are stable across the life course in Drosophila simulans.

Within populations, adult sex ratios influence population growth and extinction risk, mating behaviours and parental care. Sex ratio adjustment can also have pronounced effects on individual fitness. Accordingly, it is important that we understand how often, and why, offspring sex ratios deviate from parity. In Drosophila melanogaster, females appear to improve their fitness by producing fewer sons when paired with older males. However, facultative sex ratio adjustment in D. melanogaster is controversial, and our understanding of how sex ratio skew affects fitness is hampered by pronounced sexual conflict in this species. Additionally, it is unclear whether maternal age or quality interacts with paternal age to influence offspring sex ratios. Here, we test whether offspring sex ratios vary as a function of maternal quality, and maternal and paternal age in Drosophila simulans, a sister species of D. melanogaster that lacks overt sexual conflict. We find that offspring sex ratios are slightly male-biased overall, but constant across the female life course, and independent of female quality, or paternal age. To really understand if, how and when females skew offspring sex ratios, we need studies linking offspring sex ratios to paternal and maternal phenotypes that are predicted to shift optimal investment in sons and daughters.

Selection on age of female reproduction in the marula fruit fly, Ceratitis cosyra (Walker) (Diptera: Tephritidae), decreases total antioxidant capacity and lipid peroxidation.

The oxidative damage caused to cells by Reactive Oxygen Species (ROS) is one of several factors implicated in causing ageing. Oxidative damage may also be a proximate cost of reproductive effort that mediates the trade-off often observed between reproduction and survival. However, how the balance between oxidative damage and antioxidant protection affects life-history strategies is not fully understood. To improve our understanding, we selected on female reproductive age in the marula fruit fly, Ceratitis cosyra, and quantified the impact of selection on female and male mortality risk, female fecundity, male sperm transfer, calling and mating. Against expectations, upward-selected lines lived shorter lives and experienced some reductions in reproductive performance. Selection affected oxidative damage to lipids and total antioxidant protection, but not in the direction predicted; longer lives were associated with elevated oxidative damage, arguing against the idea that accumulated oxidative damage reduces lifespan. Greater reproductive effort was also associated with elevated oxidative damage, suggesting that oxidative damage may be a cost of reproduction, although one that did not affect survival. Our results add to a body of data showing that the relationship between lifespan, reproduction and oxidative damage is more complex than predicted by existing theories.

Exploring the role of host specialisation and oxidative stress in interspecific lifespan variation in subtropical tephritid flies.

In herbivorous insects, the degree of host specialisation may be one ecological factor that shapes lifespan. Because host specialists can only exploit a limited number of plants, their lifecycle should be synchronised with host phenology to allow reproduction when suitable hosts are available. For species not undergoing diapause or dormancy, one strategy to achieve this could be evolving long lifespans. From a physiological perspective, oxidative stress could explain how lifespan is related to degree of host specialisation. Oxidative stress caused by Reactive Oxygen Species (ROS) might help underpin ageing (the Free Radical Theory of Aging (FRTA)) and mediate differences in lifespan. Here, we investigated how lifespan is shaped by the degree of host specialisation, phylogeny, oxidative damage accumulation and antioxidant protection in eight species of true fruit flies (Diptera: Tephritidae). We found that lifespan was not constrained by species relatedness or oxidative damage (arguing against the FRTA); nevertheless, average lifespan was positively associated with antioxidant protection. There was no lifespan difference between generalist and specialist species, but most of the tephritids studied had long lifespans in comparison with other dipterans. Long lifespan may be a trait under selection in fruit-feeding insects that do not use diapause.

Complex interactions between sperm viability and female fertility.

Sperm viability is a major male fitness component, with higher sperm viability associated with enhanced sperm competitiveness. While many studies have focussed on sperm viability from the male fitness standpoint, its impact on female fitness is less clear. Here we used a panel of 32 isogenic Drosophila simulans lines to test for genetic variation in sperm viability (percentage of viable cells). We then tested whether sperm viability affected female fitness by mating females to males from low or high sperm viability genotypes. We found significant variation in sperm viability among genotypes, and consistent with this, sperm viability was highly repeatable within genotypes. Additionally, females mated to high sperm viability males laid more eggs in the first seven hours after mating, and produced more offspring in total. However, the early increase in oviposition did not result in more offspring in the 8 hours following mating, suggesting that mating with high sperm-viability genotypes leads to egg wastage for females shortly after copulation. Although mating with high sperm-viability males resulted in higher female fitness in the long term, high quality ejaculates would result in a short-term female fitness penalty, or at least lower realised fitness, potentially generating sexual conflict over optimal sperm viability.

Prioritising physical and psychological symptoms: what are the barriers and facilitators to the discussion of anxiety in the primary care consultation?

BACKGROUND: Anxiety is under-recorded and under-treated in the UK and is under-represented in research compared with depression. Detecting anxiety can be difficult because of co-existing conditions. GPs can be reluctant to medicalise anxiety symptoms and patients can be reluctant to disclose them, for a variety of reasons. This research addresses the gap in evidence of real-life consultations of patients with anxiety and explores how physical and psychological symptoms are discussed and prioritised by patients and GPs in primary care consultations. METHODS: A mixed methods study using a baseline questionnaire, video-recorded primary care consultations and interview data with patients and GPs. RESULTS: Seventeen patients with anxiety symptoms (GAD-7 score ≥ 10) completed a questionnaire, had their consultation video-recorded and took part in a semi-structured interview. Four GPs were interviewed. The main themes that emerged from GP and patients accounts as barriers and facilitators to discussing anxiety mostly mirrored each other. The GP/patient relationship and continuity of care was the main facilitator for the discussion of anxiety in the consultation. The main barriers were: attribution of or unacknowledged symptoms; co-morbidities; and time constraints. GPs overcame these barriers by making repeat appointments and employing prioritising techniques; patients by choosing an empathetic GP. CONCLUSIONS: The findings add to the evidence base concerning the management of anxiety in primary care. The findings suggest that the discussion around anxiety is a process negotiated between the patient and the GP influenced by a range of barriers and facilitators. Co-existing depression and health anxieties can mask anxiety symptoms in patients. Good practice techniques such as bringing back patients for appointments to foster continuity of care and understanding can help disclosure and detection of anxiety symptoms. Future research could investigate this longitudinally and should include a wider range of GPs practices and GPs.

Sexual conflict.

Evolutionary conflict arises from differences in the fitness interests of replicating entities and has its roots in relatedness asymmetries. Every replicator is related to itself by 100%, but in most cases is less related to other replicators, which generates selfishness and conflicts of interest. Since this basic condition is the norm at many levels of biological organization, conflict is rife in biological systems. Sexual conflict, on which we focus here, is the evolutionary conflict that occurs between males and females because of their divergent fitness interests. Sexual conflict occurs despite sexual reproduction requiring some level of cooperation between males and females because the fitness interests of the sexes are nevertheless never perfectly aligned. In other words, males and females may agree on where they are going, but not necessarily on how to get there. Sexual conflict is a vast topic with relevance to many areas of biology and so here we restrict our focus to matters we think are of broadest interest.

Extracellular Vesicles in the Synovial Joint: Is there a Role in the Pathophysiology of Osteoarthritis?

The role of extracellular vesicles (EV) in osteoarthritis has become the focus of much research. These vesicles were isolated from several cell types found in synovial joint including chondrocytes and synovium. As articular cartilage is an avascular tissue surrounded by synovial fluid, it is believed that EV might play a crucial role in the homeostasis of cartilage and also could hold key information in the pathogenesis of osteoarthritis. This is thought to be due to activation of pro-inflammatory factors leading to a catabolic state and degradation of cartilage. In addition, due to the nature of articular cartilage lacking neuronal innervation, knowledge of EV can contribute to identification of novel biomarkers in this debilitating condition. This can be either directly isolated from aspirate of synovial fluid or from peripheral blood. Finally, EVs are known to shuttle important signalling molecules which can be utilised as unique modality in transferring therapeutic compounds in a cell free manner.

Wolbachia infection can bias estimates of intralocus sexual conflict.

Males and females share most of their genome and develop many of the same traits. However, each sex frequently has different optimal values for these shared traits, creating intralocus sexual conflict. This conflict has been observed in wild and laboratory populations of insects and affects important evolutionary processes such as sexual selection, the maintenance of genetic variation, and possibly even speciation. Given the broad impacts of intralocus conflict, accurately detecting and measuring it is important. A common way to detect intralocus sexual conflict is to calculate the intersexual genetic correlation for fitness, with negative values suggesting conflict. Here, we highlight a potential confounder of this measure-cytoplasmic incompatibility caused by the intracellular parasite Wolbachia. Infection with Wolbachia can generate negative intersexual genetic correlations for fitness in insects, suggestive of intralocus sexual conflict. This is because cytoplasmic incompatibility reduces the fitness of uninfected females mated to infected males, while uninfected males will not suffer reductions in fitness if they mate with infected females and may even be fitter than infected males. This can lead to strong negative intersexual genetic correlations for fitness, mimicking intralocus conflict. We illustrate this issue using simulations and then present Drosophila simulans data that show how reproductive incompatibilities caused by Wolbachia infection can generate signals of intralocus sexual conflict. Given that Wolbachia infection in insect populations is pervasive, but populations usually contain both infected and uninfected individuals providing scope for cytoplasmic incompatibility, this is an important consideration for sexual conflict research but one which, to date, has been largely underappreciated.

Penis evolution across species: divergence and diversity.

The penis is an incredibly diverse and rapidly evolving structure, such that even in closely related species that otherwise differ very little in their morphology, penis form can be highly differentiated. Penises are also much more complex than their fundamental function - sperm transfer - would seem to require. The rapid divergent evolution of male structures is typically the signature of traits under sexual selection and the current evidence suggests the penis is no different in this regard. Despite the general agreement that sexual selection is the main driver of penis evolution, many questions about penis evolution remain unresolved. Furthermore, the penis might be an ideal characteristic on which to focus in the drive to link phenotype with genotype.