Cost of reproduction in the Queensland fruit fly: Y-model versus lethal protein hypothesis

The trade-off between lifespan and reproduction is commonly explained by differential allocation of limited resources. Recent research has shown that the ratio of protein to carbohydrate (P : C) of a fly''s diet mediates the lifespan–reproduction trade-off, with higher P : C diets increasing egg production but decreasing lifespan. To test whether this P : C effect is because of changing allocation strategies (Y-model hypothesis) or detrimental effects of protein ingestion on lifespan (lethal protein hypothesis), we measured lifespan and egg production in Queensland fruit flies varying in reproductive status (mated, virgin and sterilized females, virgin males) that were fed one of 18 diets varying in protein and carbohydrate amounts. The Y-model predicts that for sterilized females and for males, which require little protein for reproduction, there will be no effect of P : C ratio on lifespan; the lethal protein hypothesis predicts that the effect of P : C ratio should be similar in all groups. In support of the lethal protein hypothesis, and counter to the Y-model, the P : C ratio of the ingested diets had similar effects for all groups. We conclude that the trade-off between lifespan and reproduction is mediated by the detrimental side-effects of protein ingestion on lifespan.     

Dietary choice for a balanced nutrient intake increases the mean and reduces the variance in the reproductive performance of male and female cockroaches

Sexual selection may cause dietary requirements for reproduction to diverge across the sexes and promote the evolution of different foraging strategies in males and females. However, our understanding of how the sexes regulate their nutrition and the effects that this has on sex‐specific fitness is limited. We quantified how protein (P) and carbohydrate (C) intakes affect reproductive traits in male (pheromone expression) and female (clutch size and gestation time) cockroaches (Nauphoeta cinerea). We then determined how the sexes regulate their intake of nutrients when restricted to a single diet and when given dietary choice and how this affected expression of these important reproductive traits. Pheromone levels that improve male attractiveness, female clutch size and gestation time all peaked at a high daily intake of P:C in a 1:8 ratio. This is surprising because female insects typically require more P than males to maximize reproduction. The relatively low P requirement of females may reflect the action of cockroach endosymbionts that help recycle stored nitrogen for protein synthesis. When constrained to a single diet, both sexes prioritized regulating their daily intake of P over C, although this prioritization was stronger in females than males. When given the choice between diets, both sexes actively regulated their intake of nutrients at a 1:4.8 P:C ratio. The P:C ratio did not overlap exactly with the intake of nutrients that optimized reproductive trait expression. Despite this, cockroaches of both sexes that were given dietary choice generally improved the mean and reduced the variance in all reproductive traits we measured relative to animals fed a single diet from the diet choice pair. This pattern was not as strong when compared to the single best diet in our geometric array, suggesting that the relationship between nutrient balancing and reproduction is complex in this species.     

Effects of macronutrient intake on the lifespan and fecundity of the marula fruit fly,Ceratitis cosyra (Tephritidae): Extreme lifespan in a host specialist

In insects, lifespan and reproduction are strongly associated with nutrition. The ratio and amount of nutrients individuals consume affect their life expectancy and reproductive investment. The geometric framework (GF) enables us to explore how animals regulate their intake of multiple nutrients simultaneously and determine how these nutrients interact to affect life‐history traits of interest. Studies using the GF on host‐generalist tephritid flies have highlighted trade‐offs between longevity and reproductive effort in females, mediated by the protein‐to‐carbohydrate (P:C) ratio that individuals consume. Here, we tested how P and C intake affect lifespan (LS) in both sexes, and female lifetime (LEP), and daily (DEP) egg production, in Ceratitis cosyra, a host‐specialist tephritid fly. We then determined the P:C ratio that C. cosyra defends when offered a choice of foods. Female LS was optimized at a 0:1 P:C ratio, whereas to maximize their fecundity, females needed to consume a higher P:C ratio (LEP = 1:6 P:C; DEP = 1:2.5 P:C). In males, LS was also optimized at a low P:C ratio of 1:10. However, when given the opportunity to regulate their intake, both sexes actively defended a 1:3 P:C ratio, which is closer to the target for DEP than either LS or LEP. Our results show that female C. cosyra experienced a moderate trade‐off between LS and fecundity. Moreover, the diets that maximized expression of LEP and DEP were of lower P:C ratio than those required for optimal expression of these traits in host‐generalist tephritids or other generalist insects.     

Sex‐specific effects of protein and carbohydrate intake on reproduction but not lifespan in Drosophila melanogaster

Modest dietary restriction extends lifespan (LS) in a diverse range of taxa and typically has a larger effect in females than males. Traditionally, this has been attributed to a stronger trade‐off between LS and reproduction in females than in males that is mediated by the intake of calories. Recent studies, however, suggest that it is the intake of specific nutrients that extends LS and mediates this trade‐off. Here, we used the geometric framework (GF) to examine the sex‐specific effects of protein (P) and carbohydrate (C) intake on LS and reproduction in Drosophila melanogaster. We found that LS was maximized at a high intake of C and a low intake of P in both sexes, whereas nutrient intake had divergent effects on reproduction. Male offspring production rate and LS were maximized at the same intake of nutrients, whereas female egg production rate was maximized at a high intake of diets with a P:C ratio of 1:2. This resulted in larger differences in nutrient‐dependent optima for LS and reproduction in females than in males, as well as an optimal intake of nutrients for lifetime reproduction that differed between the sexes. Under dietary choice, the sexes followed similar feeding trajectories regulated around a P:C ratio of 1:4. Consequently, neither sex reached their nutritional optimum for lifetime reproduction, suggesting intralocus sexual conflict over nutrient optimization. Our study shows clear sex differences in the nutritional requirements of reproduction in D. melanogaster and joins the growing list of studies challenging the role of caloric restriction in extending LS.     

Dietary protein:carbohydrate balance is a critical modulator of lifespan and reproduction in Drosophila melanogaster: A test using a chemically defined diet

Macronutrient balance is an important determinant of fitness in many animals, including insects. Previous studies have shown that altering the concentrations of yeast and sugar in the semi-synthetic media has a profound impact on lifespan in Drosophila melanogaster, suggesting that dietary protein:carbohydrate (P:C) balance is the main driver of lifespan and ageing processes. However, since yeast is rich in multiple nutrients other than proteins, this lifespan-determining role of dietary P:C balance needs to be further substantiated through trials using a chemically-defined, synthetic diet. In the present study, the effects of dietary P:C balance on lifespan and fecundity were investigated in female D. melanogaster flies fed on one of eight isocaloric synthetic diets differing in P:C ratio (0:1, 1:16, 1:8, 1:4, 1:2, 1:1, 2:1 or 4:1). Lifespan and dietary P:C ratio were related in a convex manner, with lifespan increasing to a peak at the two intermediate P:C ratios (1:2 and 1:4) and falling at the imbalanced ratios (0:1 and 4:1). Ingesting nutritionally imbalanced diets not only caused an earlier onset of senescence but also accelerated the age-dependent increase in mortality. Egg production was suppressed when flies were fed on a protein-deficient food (0:1), but increased with increasing dietary P:C ratio. Long-lived flies at the intermediate P:C ratios (1:2 and 1:4) stored a greater amount of lipids than those short-lived ones at the two imbalanced ratios (0:1 and 4:1). These findings provide a strong support to the notion that adequate dietary P:C balance is crucial for extending lifespan in D. melanogaster and offer new insights into how dietary P:C balance affects lifespan and ageing through its impacts on body composition.     

Female access and diet affect insemination success,senescence and the cost of reproduction in the male Mexican fruit fly Anastrepha ludens

Hypotheses exploring the influence of dietary conditions on the life‐history trade‐off between survival and reproductive success are extensively tested in female insects but only rarely explored in males. The present study examines the impact of dietary quality and female access on age‐specific reproduction and survival of the male Mexican fruit fly Anastrepha ludens Loew (Diptera: Tephritidae). There is a clear cost of female access for males with access to dietary protein, measurable as a decrease in life expectancy, which is further influenced by the age when females are introduced. A protein deficient diet reduces the lifespan benefit of virginity and masks the detrimental effect of female access on male life expectancy. Dietary protein is not necessary for reproductive success, although access to protein at eclosion improves the lifetime reproductive success of males compared to when it is delayed. Overall, reproductive success diminishes as the male flies age, regardless of the dietary conditions, providing evidence for reproductive senescence in males. Delaying the males' access to a protein source fails to influence the negative effect of age on reproductive ability. Because age‐specific reproductive rates decline with age, regardless of diet, male fitness does not benefit from lifespan extension. Therefore, males can be expected to allocate available resources towards reproductive effort in favour of an extended lifespan, regardless of mate and protein availability.     

Sex differences in nutrient intake can reduce the potential for sexual conflict over fitness maximization by female and male crickets

As females and males have different roles in reproduction, they are expected to require different nutrients for the expression of reproductive traits. However, due to their shared genome, both sexes may be constrained in the regulation of nutrient intake that maximizes sex‐specific fitness. Here, we used the Geometric Framework for nutrition to examine the effect of macronutrient and micronutrient intakes on lifespan, fecundity and cuticular hydrocarbons (CHCs) that signal mate quality to prospective mates in female field crickets, Teleogryllus oceanicus. In addition, we contrasted nutritional effects on life‐history traits between males and females to determine how sex differences influence nutrient regulation. We found that carbohydrate intake maximized female lifespan and protein intake influenced CHC expression, while early life fecundity (cumulative fecundity at day 21) and lifetime fecundity were dependent on both macronutrient and micronutrient intakes. Fecundity required different nutrient blends to those required to optimize sperm viability in males, generating the potential for sexual conflict over macronutrient intake. The regulation of protein (P) and carbohydrate (C) intakes by virgin and mated females initially matched that of males, but females adjusted their intake to a higher P:C ratio, 1P:2C, that maximized fecundity as they aged. This suggests that a sex‐specific, age‐dependent change in intake target for sexually mature females, regardless of their mating status, adjusts protein consumption in preparation for oviposition. Sex differences in the regulation of nutrient intake to optimize critical reproductive traits in female and male T. oceanicus provide an example of how sexual conflict over nutrition can shape differences in foraging between the sexes.     

Fitness advantage from nuptial gifts in female fireflies

Abstract 1. In many insects, males provide nuptial gifts to females in the form of spermatophores, sperm-containing structures produced by male accessory glands.
2. The work reported here examined the influence of both spermatophore number and spermatophore size on female reproductive output in two related firefly beetles, Photinus ignitus and Ellychnia corrusca (Coleoptera: Lampyridae). Based on differences in adult diet, male spermatophores were predicted to increase female reproductive output to a greater extent in P. ignitus than in E. corrusca .
3. Female fecundity was significantly higher in triply mated females than in singly mated females in both species, with no difference between mating treatments in female lifespan or egg hatching success. No effects of second male spermatophore size on fecundity, lifespan, or egg hatching success were detected in either species.
4. These results suggest a direct fitness advantage from multiple mating for females in both species, although enhanced fecundity may be due either to allocation of spermatophore nutrients to eggs or to other substances transferred within the spermatophore acting as oviposition stimulants.     

Towards a synthesis of frameworks in nutritional ecology: interacting effects of protein,carbohydrate and phosphorus on field cricket fitness

Phosphorus has been identified as an important determinant of nutrition-related biological variation. The macronutrients protein (P) and carbohydrates (C), both alone and interactively, are known to affect animal performance. No study, however, has investigated the importance of phosphorus relative to dietary protein or carbohydrates, or the interactive effects of phosphorus with these macronutrients, on fitness-related traits in animals. We used a nutritional geometry framework to address this question in adult field crickets (Gryllus veletis). Our results showed that lifespan, weight gain, acoustic mate signalling and egg production were maximized on diets with different P : C ratios, that phosphorus did not positively affect any of these fitness traits, and that males and females had different optimal macronutrient intake ratios for reproductive performance. When given a choice, crickets selected diets that maximized both lifespan and reproductive performance by preferentially eating diets with low P : C ratios, and females selected diets with a higher P : C ratio than males. Conversely, phosphorus intake was not regulated. Overall, our findings highlight the importance of disentangling the influences of different nutrients, and of quantifying both their individual and interactive effects, on animal fitness traits, so as to gain a more integrative understanding of their nutritional ecology.     

Effects of diet and host access on fecundity and lifespan in two fruit fly species with different life‐history patterns

The reproductive ability of female tephritids can be limited and prevented by denying access to host plants and restricting the dietary precursors of vitellogenesis. The mechanisms underlying the delayed egg production in each case are initiated by different physiological processes that are anticipated to have dissimilar effects on lifespan and reproductive ability later in life. The egg‐laying abilities of laboratory‐reared females of the Mediterranean fruit fly (Ceratitis capitata Wiedmann) and melon fly (Bactrocera cucurbitae Coquillett) from Hawaii are delayed or suppressed by limiting access to host fruits and dietary protein. In each case, this is expected to prevent the loss of lifespan associated with reproduction until protein or hosts are introduced. Two trends are observed in each species: first, access to protein at eclosion leads to a greater probability of survival and a higher reproductive ability than if it is delayed and, second, delayed host access reduces lifetime reproductive ability without improving life expectancy. When host access and protein availability are delayed, the rate of reproductive senescence is reduced in the medfly, whereas the rate of reproductive senescence is generally increased in the melon fly. Overall, delaying reproduction lowers the fitness of females by constraining their fecundity for the remainder of the lifespan without extending the lifespan. © 2013 The Royal Entomological Society     

Adult dietary protein has age‐ and context‐dependent effects on male post‐copulatory performance

The highly conserved effect of dietary protein restriction on lifespan and ageing is observed in both sexes and across a vast range of taxa. This extension of lifespan is frequently accompanied by a reduction in female fecundity, and it has been hypothesized that individuals may reallocate resources away from reproduction and into somatic maintenance. However, effects of dietary protein restriction on male reproduction are less consistent, suggesting that these effects may depend on other environmental parameters. Using the neriid fly, Telostylinus angusticollis, we examined age‐specific effects of adult dietary protein restriction on male post‐copulatory reproductive performance (fecundity and offspring viability). To explore the context dependence of these effects, we simultaneously manipulated male larval diet and adult mating history. We found that protein‐restricted males sired less viable offspring at young ages, but offspring viability increased with paternal age and eventually exceeded that of fully fed males. The number of eggs laid by females was not affected by male dietary protein, whereas egg hatching success was subject to a complex interaction of male adult diet, age, larval diet and mating history. These findings suggest that effects of protein restriction on male reproduction are highly context dependent and cannot be explained by a simple reallocation of resources from reproduction to somatic maintenance. Rather, these effects appear to involve changes in the scheduling of male reproductive investment with age.     

Nutrients, not caloric restriction, extend lifespan in Queensland fruit flies (Bactrocera tryoni)

Caloric restriction (CR) has been widely accepted as a mechanism explaining increased lifespan (LS) in organisms subjected to dietary restriction (DR), but recent studies investigating the role of nutrients have challenged the role of CR in extending longevity. Fuelling this debate is the difficulty in experimentally disentangling CR and nutrient effects due to compensatory feeding (CF) behaviour. We quantified CF by measuring the volume of solution imbibed and determined how calories and nutrients influenced LS and fecundity in unmated females of the Queensland fruit fly, Bactocera tryoni (Diptera: Tephritidae). We restricted flies to one of 28 diets varying in carbohydrate:protein (C:P) ratios and concentrations. On imbalanced diets, flies overcame dietary dilutions, consuming similar caloric intakes for most dilutions. The response surface for LS revealed that increasing C:P ratio while keeping calories constant extended LS, with the maximum LS along C:P ratio of 21:1. In general, LS was reduced as caloric intake decreased. Lifetime egg production was maximized at a C:P ratio of 3:1. When given a choice of separate sucrose and yeast solutions, each at one of five concentrations (yielding 25 choice treatments), flies regulated their nutrient intake to match C:P ratio of 3:1. Our results (i) demonstrate that CF can overcome dietary dilutions; (ii) reveal difficulties with methods presenting fixed amounts of liquid diet; (iii) illustrate the need to measure intake to account for CF in DR studies and (iv) highlight nutrients rather than CR as a dominant influence on LS.     

Ant workers die young and colonies collapse when fed a high-protein diet

A key determinant of the relationship between diet and longevity is the balance of protein and carbohydrate in the diet. Eating excess protein relative to carbohydrate shortens lifespan in solitary insects. Here, we investigated the link between high-protein diet and longevity, both at the level of individual ants and colonies in black garden ants, Lasius niger. We explored how lifespan was affected by the dietary protein-to-carbohydrate ratio and the duration of exposure to a high-protein diet. We show that (i) restriction to high-protein, low-carbohydrate diets decreased worker lifespan by up to 10-fold; (ii) reduction in lifespan on such diets was mainly due to elevated intake of protein rather than lack of carbohydrate; and (iii) only one day of exposure to a high-protein diet had dire consequences for workers and the colony, reducing population size by more than 20 per cent.     

Resource allocation to reproduction and soma in Drosophila: a stable isotope analysis of carbon from dietary sugar

Metabolic resources in adults of holometabolous insects may derive either from larval or adult feeding. In Drosophila melanogaster, reproduction and lifespan are differently affected by larval vs. adult resource availability, and it is unknown how larval vs. adult acquired nutrients are differentially allocated to somatic and reproductive function. Here we describe the allocation of carbon derived from dietary sugar in aging female D. melanogaster. Larval and adult flies were fed diets contrasting in sucrose (13)C/(12)C, from which we determined the extent to which carbon acquired at each stage contributed to adult somatic tissue and to egg manufacture. Dietary sugar is very important in egg provisioning; at every age, roughly one half of the carbon in eggs was derived from sugar, which turned over from predominantly larval to entirely adult dietary sources. Sucrose provided approximately 40% of total somatic carbon, of which adult dietary sucrose came to supply approximately 75%. Unlike in eggs, however, adult acquired sucrose did not entirely replace the somatic carbon from larvally acquired sucrose. Because carbon from larval sucrose appears to be fairly "replaceable", larval sucrose cannot be a limiting substrate in resource allocation between reproduction and lifespan.     

Sexes suffer from suboptimal lifespan because of genetic conflict in a seed beetle

Males and females have different routes to successful reproduction, resulting in sex differences in lifespan and age-specific allocation of reproductive effort. The trade-off between current and future reproduction is often resolved differently by males and females, and both sexes can be constrained in their ability to reach their sex-specific optima owing to intralocus sexual conflict. Such genetic antagonism may have profound implications for evolution, but its role in ageing and lifespan remains unresolved. We provide direct experimental evidence that males live longer and females live shorter than necessary to maximize their relative fitness in Callosobruchus maculatus seed beetles. Using artificial selection in a genetically heterogeneous population, we created replicate long-life lines where males lived on average 27 per cent longer than in short-life lines. As predicted by theory, subsequent assays revealed that upward selection on male lifespan decreased relative male fitness but increased relative female fitness compared with downward selection. Thus, we demonstrate that lifespan-extending genes can help one sex while harming the other. Our results show that sexual antagonism constrains adaptive life-history evolution, support a novel way of maintaining genetic variation for lifespan and argue for better integration of sex effects into applied research programmes aimed at lifespan extension.     

Sex-specific fitness effects of nutrient intake on reproduction and lifespan

Diet affects both lifespan and reproduction [1-9], leading to the prediction that the contrasting reproductive strategies of the sexes should result in sex-specific effects of nutrition on fitness and longevity [6, 10] and favor different patterns of nutrient intake in males and females. However, males and females share most of their genome and intralocus sexual conflict may prevent sex-specific diet optimization. We show that both male and female longevity were maximized on a high-carbohydrate low-protein diet in field crickets Teleogryllus commodus, but male and female lifetime reproductive performances were maximized in markedly different parts of the nutrient intake landscape. Given a choice, crickets exhibited sex-specific dietary preference in the direction that increases reproductive performance, but this sexual dimorphism in preference was incomplete, with both sexes displaced from the optimum diet for lifetime reproduction. Sexes are, therefore, constrained in their ability to reach their sex-specific dietary optima by the shared biology of diet choice. Our data suggest that sex-specific selection has thus far failed fully to resolve intralocus sexual conflict over diet optimization. Such conflict may be an important factor linking nutrition and reproduction to lifespan and aging.     

Effects of dietary protein:carbohydrate balance on life-history traits in six laboratory strains of Drosophila melanogaster

Drosophila melanogaster Meigen (Diptera: Drosophilidae) is a key model insect for studying life span and aging. Many laboratory strains of D. melanogaster are currently used by laboratories worldwide, but they are known to vary considerably in their physiology, behavior, and life histories. Although the importance of dietary protein:carbohydrate (P:C) balance as a predominant determinant of life span and other life-history traits has been highlighted in recent research, it remains unexplored whether the impacts of P:C balance on these fitness-related traits vary in a strain-specific manner in D. melanogaster. In this study, we compared the life-history consequences (life span, egg production rate, pre-adult survival, development time, and body mass at eclosion) of six laboratory strains of D. melanogaster (w1118, yw, Oregon-R, white Canton-S, Canton-S-SNU, and Canton-S-Inha) allocated to one of four synthetic diets differing in P:C ratio (1:16, 1:4, 1:1, or 4:1). The effects of dietary P:C balance on various adult and larval life-history traits were qualitatively similar across all strains studied in this study. Regardless of fly strain, adults exhibited a shortened life span and improved egg production on a diet with the highest P:C ratio of 4:1. In all strains, larvae raised on a diet comprising the lowest P:C ratio of 1:16 suffered high mortality, delayed development time, and reduced body mass. Despite the general similarity in the direction of the effect of P:C balance across strains, fly strains differed in the magnitude of their life-history responses to dietary P:C balance, as indicated by a significant interaction between fly strain and dietary P:C ratio for all measured traits except body mass at eclosion. Possible mechanisms explaining such strain-specific responses are discussed.     

Diet, sex, and death in field crickets

Senescence is shaped by age-dependent trade-offs between fitness components. Because males and females invest different resources in reproduction, the trade-offs behind age-dependent reproductive effort should be resolved differently in the sexes. In this study, we assess the effects of diet (high carbohydrate and low protein vs. equal carbohydrate and protein) and mating (once mated vs. virgin) on lifespan and age-dependent mortality in male and female field crickets (Teleogryllus commodus), and on male calling effort. Females always had higher actuarial ageing rates than males, and we found a clear lifespan cost of mating in females. Mated males, however, lived longer than virgin males, possibly because virgins call more than mated males. The fastest age-dependent increases in mortality were among mated males on the high-carbohydrate diet. Males on a high-carbohydrate diet showed a faster increase in calling effort earlier in life, and a more pronounced pattern of senescence once they reached this peak than did males on a diet with equal amounts of protein and carbohydrates. Our results provide evidence that the cost of mating in this cricket species is both diet and sex-dependent, and that the underlying causes of sex differences in life-history traits such as lifespan and senescence can be complex.     

Differences in the pre‐diapause and pre‐oviposition accumulation of critical nutrients in adult females of the beetle Colaphellus bowringi

In many insect species, the differentiation of development between diapause and reproduction first becomes obvious during the diapause preparation (pre‐diapause) and pre‐oviposition phases. However, the differentiation of nutrient accumulation between these two phases remains unclear. We compared the weights of pre‐diapause and reproductive adult female Colaphellus bowringi Baly (Coleoptera: Chrysomelidae), and measured their triacylglycerol (TAG), protein, and carbohydrate content from emergence until they had fed for 4 days post‐eclosion. We also compared the ovarian development and accumulation of lipid droplets between pre‐diapause and reproductive adult females in order to determine whether we could visually detect differences in nutrient allocation. The weights of both pre‐diapause and reproductive females increased with duration of feeding. The fresh weight and water content of pre‐diapause females was significantly lower than that of reproductive females after feeding for 3 days post‐eclosion. Pre‐diapause females channeled their reserves into TAG in the fat body, whereas reproductive females converted nutrients into proteins and carbohydrates for egg development. These results quantify differences in nutrient accumulation between pre‐diapause and reproductive adult female C. bowringi, and provide clues for understanding the molecular mechanisms underlying differences in the allocation of nutrients between diapause and reproduction in insects.     

Effects of adult nutrition on female reproduction in a fruit-feeding butterfly: the role of fruit decay and dietary lipids

It was generally believed that butterflies and other holometabolous insects rely primarily on reserves accumulated during the larval stage for reproduction. Recent studies, however, highlight the often fundamental importance of adult nutrition to realize the full reproductive potential. While the importance of carbohydrates is fairly well understood, the role of most other adult-derived substances is only partially resolved. We here focus on the effects of dietary lipids (cholesterol, polyunsaturated fatty acids) and fruit decay (dietary yeast, ethanol) on female reproduction in the tropical, fruit-feeding butterfly Bicyclus anynana (Nymphalidae). We found that banana-fed control females outperformed all other groups fed on sucrose-based diets. Lipids, yeast or ethanol added to a sugar solution did not yield a similarly high reproductive output compared to fruit-fed females. Groups fed fresh or decaying banana showed no differences in reproductive performance. As we could not identify a single pivotal substance, we conclude that resource congruence (the use of nutrient types in a specified ratio) rather than any specific nutrient component is of key importance for maximum reproductive output. Further, dietary quality may affect egg hatching success in spite of no obvious effects on egg size and number. Thus, any implications about potential fitness effects of different diets need to consider egg (and hatchling) viability in addition to fecundity.