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Kim Jensen

Aarhus University, Department of Animal and Veterinary Sciences, Adjunct

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Phone: +45 93508121
Address: Aarhus University
Department of Animal and Veterinary Sciences
Monogastric Nutrition
Blichers Allé 20
8830 Tjele
Denmark

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Papers by Kim Jensen

The geometry of nutrient space-based life-history trade-offs: sex-specific effects of macronutrient intake on the trade-off between encapsulation ability and reproductive effort in decorated crickets

American Naturalist, 2018

Life-history theory assumes that traits compete for limited resources, resulting in trade-offs. T... more Life-history theory assumes that traits compete for limited resources, resulting in trade-offs. The most commonly manipulated resource in empirical studies is the quantity or quality of diet. Recent studies using the geometric framework for nutrition, however, suggest that trade-offs are often regulated by the intake of specific nutrients, but a formal approach to identify and quantify the strength of such trade-offs is lacking. We posit that trade-offs occur whenever life-history traits are maximized in different regions of nutrient space, as evidenced by nonoverlapping 95% confidence regions of the global maximum for each trait and large angles (v) between linear nutritional vectors and Euclidean distances (d) between global maxima. We then examined the effects of protein and carbohydrate intake on the tradeoff between reproduction and aspects of immune function in male and female Gryllodes sigillatus. Female encapsulation ability and egg production increased with the intake of both nutrients, whereas male encapsulation ability increased with protein intake but calling effort increased with carbohydrate intake. The trade-offs between traits was therefore larger in males than in females, as demonstrated by significant negative correlations between the traits in males, nonoverlapping 95% confidence regions, and larger estimates of v and d. Under dietary choice, the sexes had similar regulated intakes, but neither optimally regulated nutrient intake for maximal trait expression. We highlight the fact that greater consideration of specific nutrient intake is needed when examining nutrient space-based trade-offs.

Intralocus sexual conflict over optimal nutrient intake and the evolution of sex differences in life span and reproduction

Functional Ecology, 2022

1. Despite widespread variation in life span across species, three clear patterns exist: sex diff... more 1. Despite widespread variation in life span across species, three clear patterns exist: sex differences in life span are ubiquitous, life span is commonly traded against reproduction, and nutrition has a major influence on these traits and how they trade-off. One process that potentially unites these patterns is intralocus sexual conflict over the optimal intake of nutrients for life span and reproduction. If nutrient intake has sex-specific effects on life span and reproduction but nutrient choice is genetically linked across the sexes, intralocus sexual conflict will occur and may prevent one or both sexes from feeding to their nutritional optima.
2. Here we determine the potential for this process to operate in the decorated cricket Gryllodes sigillatus. Using the Geometric Framework for Nutrition, we restrict male and female crickets to diets varying in the ratio of protein to carbohydrates and total nutrient content to quantify the effects on life span and daily reproductive effort in the sexes. We then use inbred lines to estimate the quantitative genetic basis of nutrient choice in males and females. We combine the nutrient effects and genetic estimates to predict the magnitude of evolutionary constraint for these traits in each sex. Finally, we present male and female crickets with a much broader range of diet pairs to determine how the sexes actively regulate their intake of nutrients.
3. We show that protein and carbohydrate intake have contrasting effects on life span and reproduction in the sexes and that there are strong positive intersexual genetic correlations for the intake of these nutrients under dietary choice. This is predicted to accelerate the evolutionary response of nutrient intake in males but constrain it in females, suggesting they are losing the conflict. Supporting this view, males and females regulate nutrient intake to a common nutrient ratio that was not perfectly optimal for life span or reproduction in either sex, especially in females.
4. Our findings show that intralocus sexual conflict over the optimal intake of nutrients is likely to be an important process generating sex differences in life span and reproduction and may help explain why females age faster and live shorter than males in G. sigillatus.

Rapid evolutionary adaptation to diet composition in the black soldier fly (Hermetia illucens)

Insects, 2023

Genetic adaptation of Hermetia illucens (BSF) to suboptimal single sourced waste streams can open... more Genetic adaptation of Hermetia illucens (BSF) to suboptimal single sourced waste streams can open new perspectives for insect production. Here, four BSF lines were maintained on a single sourced, low-quality wheat bran diet (WB) or on a high-quality chicken feed diet (CF) for 13 generations. We continuously evaluated presumed evolutionary responses in several performance traits to rearing on the two diets. Subsequently, we tested responses to interchanged diets, i.e., of larvae that had been reared on low-quality feed and tested on high-quality feed and vice versa to evaluate costs associated with adaptation to different diets. BSF were found to experience rapid adaptation to the diet composition. While performances on the WB diet were always inferior to the CF diet, the adaptive responses were stronger to the former diet. This stronger response was likely due to stronger selection pressure experienced by BSF fed on the low-quality single sourced diet. The interchanged diet experiment found no costs associated with diet adaptation, but revealed cross generational gain associated with the parental CF diet treatment. Our results revealed that BSF can rapidly respond adaptively to diet, although the mechanisms are yet to be determined. This has potential to be utilized in commercial insect breeding to produce lines tailored to specific diets.

Pilot scale production of Hermetia illucens (L.) larvae and frass using former foodstuffs

Cleaner Engineering and Technology, 2022

The food and feed sector requires new sustainable sources of protein and innovative solutions for... more The food and feed sector requires new sustainable sources of protein and innovative solutions for upcycling of food waste (former foodstuffs), which today is downcycled into energy or even wasted. This study aimed at evaluating the use of former foodstuff waste streams as feed substrate for Hermetia illucens (L.) larvae (black soldier fly larvae, BSFL) under long-term and semi-industrial conditions. Different foodstuff-based mixtures and different stocking BSFL densities were used during 20 batches, and quality and safety assessments were performed on the main outputs, namely BSFL production performance, frass impurities, larval and frass nutrient profiles and heavy metal content. About 1400 kg of former foodstuffs (fresh weight) were used to produce 239 kg BSFL and 230 kg frass. The production of BSFL reared on former foodstuffs was highly efficient, with feed conversion rates (FCR) ranging between 2.3 and 5.5 (dry matter basis). The optimization experiment revealed that former foodstuffs-based mixture and high larval density (10 larvae/cm 2) lead to highly efficient (FCR: 2.6) and heavy metal-free production of BSFL and frass. The quality of the derived BSFL meal was high in terms of protein and amino acids. Furthermore, the quality of the technical frass was high in terms of N, P, and K levels and minimal packaging material residuals (<2.65%). This investigation suggests that nutrients in former foodstuffs can be successfully and safely recycled in production of BSFL.

Genotype-by-sex-by-diet interactions for nutritional preference, dietary consumption, and lipid deposition in a field cricket

Heredity, 2018

Changes in feeding behaviour, especially the overconsumption of calories, has led to a rise in th... more Changes in feeding behaviour, especially the overconsumption of calories, has led to a rise in the rates of obesity, diabetes, and other associated disorders in humans and a range of animals inhabiting human-influenced environments. However, understanding the relative contribution of genes, the nutritional environment, and their interaction to dietary intake and lipid deposition in the sexes still remains a major challenge. By combining nutritional geometry with quantitative genetics, we determined the effect of genes, the nutritional environment, and their interaction on the total nutritional preference (TP), total diet eaten (TE), and lipid mass (LM) of male and female black field crickets (Teleogryllus commodus) fed one of four diet pairs (DPs) differing in the ratio of protein to carbohydrate and total nutritional content. We found abundant additive genetic variance for TP, TE, and LM in both sexes and across all four DPs, with significant genetic correlations between TE and TP ...

Geometric Stoichiometry: unifying concepts of animal nutrition to understand how protein-rich diets can be "too much of a good thing"

Frontiers in Ecology and Evolution, 2020

Understanding the factors that control the growth of heterotrophic organisms is central to predic... more Understanding the factors that control the growth of heterotrophic organisms is central to predicting food web interactions and biogeochemical cycling within ecosystems. We present a new framework, Geometric Stoichiometry (GS), that unifies the disciplines of Nutritional Geometry (NG) and Ecological Stoichiometry (ES) by extending the equations of ES to incorporate core NG concepts, including macromolecules as currencies and the ability of animals to select foods that balance deficits and excesses of nutrients. The resulting model is used to investigate regulation of consumer growth by dietary protein:carbohydrate ratio. Growth on protein-poor diets is limited by nitrogen. Likewise, we show that growth is also diminished on protein-rich diets and that this can be mechanistically explained by means of a metabolic penalty that arises when animals use protein for energy generation. These penalties, which are incurred when dealing with the costs of producing and excreting toxic nitrogenous waste, have not hitherto been represented in standard ES theory. In order to incorporate GS within ecosystem and biogeochemical models, a new generation of integrated theoretical and experimental studies based on unified concepts of NG and ES is needed, including measurements of food selection, biomass, growth and associated physiology, and involving metabolic penalties.

Prey-specific impact of cold pre-exposure on kill rate and reproduction

Journal of Animal Ecology, 2019

1. Temperature influences biological processes of ectotherms including ecological interactions, b... more 1. Temperature influences biological processes of ectotherms including ecological interactions, but interaction strengths may depend on species-specific traits. Furthermore, ectotherms acclimate to prevailing thermal conditions by adjusting physiological parameters, which often implies costs to other fitness-related parameters. Both predators and prey may therefore pay thermal acclimation costs following exposure to suboptimal temperatures. However, these costs may be asymmetrical between predator and prey, and between the predator and different species of concurrent prey. 2. We investigated whether thermal pre-exposure affected subsequent kill rate and predator fitness when foraging on prey that differ in ease of capture, and whether changes were primarily caused by predator or by prey pre-exposure effects. Specifically, we were interested in whether there were interactions between predator pre-exposed temperature and specific prey. 3. Using the mesostigmatid mite Gaeolaelaps aculeifer as a generalist predator and the collembolans Folsomia candida and Protaphorura fimata as prey, we measured the impact of present temperature, predator pre-exposure temperature, prey pre-exposure temperature (all 10 or 20°C), prey species, and all interactions on prey numbers killed, predator eggs produced, and exploitation of killed prey in a full factorial design. 4. Mites killed P. fimata in equal numbers independent of the presence of F. candida, but killed F. candida when P. fimata was absent. Mite kill rate and reproduction were significantly affected by mite pre-exposure temperature and test temperature , but not by prey pre-exposure temperature. Significantly more of the slower prey was killed than of the quicker prey. Importantly, we found significant syner-gistic negative interaction effects between predator cold pre-exposure and hunting prey of higher agility on predator kill rate and reproduction. 5. Our findings show that the negative effects of cold and cold pre-exposure on kill rate and reproduction may be more severe when predators forage on quick prey. The study implies that predator cold exposure has consequences for specific prey survival following cold due to altered predation pressures, which in nature should influence the specific prey population dynamics and apparent competition

Increased lipid accumulation but not reduced metabolism explains improved starvation tolerance in cold-acclimated arthropod predators

Science of Nature, 2018

Predatory arthropods are used for biological control in greenhouses, but there is increasing inte... more Predatory arthropods are used for biological control in greenhouses, but there is increasing interest to extend their use to the outdoor environment where temperatures are typically lower. Acclimation at low temperature increases the ability of ectotherms to cope with subsequent more extreme cold, but may involve costs or benefits to other performance traits. A recent study in mesostigmatid mites (Gaeolaelaps aculeifer) showed that starvation tolerance was improved following a period of cold exposure. However, the physiological mechanisms that underlie improved starvation tolerance following cold exposure were not investigated. To examine whether cold acclimation would also improve starvation tolerance in an insect, we repeated the starvation study in another arthropod predator, the pirate bug Orius majusculus, as well as in G. aculeifer. Before tests, the two species were acclimated at 10, 15, or 20°C for 7 (G. aculeifer) or 16 (O. majusculus) days. We then analyzed the effects of thermal exposure on body composition, consumption, and basal metabolic rate in both species. Our results confirmed that exposure to low temperature improves starvation tolerance in these arthropod predators. Body composition analyses revealed that both species had accumulated larger lipid stores during exposure to colder temperature, which at least in part can explain the larger starvation tolerance following cold exposure. In contrast, consumption and basal metabolic rate were not changed by thermal acclimation. Our study indicates that predatory arthropods exposed to cold increase their physiological robustness and ability to endure environmental challenges, including low temperature and low prey availability.

Are commercial stocks of biological control agents genetically depauperate? -A case study on the pirate bug Orius majusculus Reuter

Biological Control, 2018

Predatory arthropods are increasingly used in biological control of insect pests. For this purpos... more Predatory arthropods are increasingly used in biological control of insect pests. For this purpose, control agents are produced commercially in large quantities for release in crops. The production stocks, however, may have undergone numerous population bottlenecks and may have been exposed to artificial selection pressures in the production facilities. Accordingly, commercial populations may be experiencing loss of genetic variation through inbreeding and genetic drift, which may reduce fitness and biocontrol efficiency. In the present study we investigated whether populations of the pirate bug Orius majusculus (Reuter) purchased from three European companies differed in a range of performance traits including predation rate, starvation tolerance, body size, locomotor activity, and heat tolerance. Furthermore, we crossed all populations pairwise and tested whether outcrossed F2 hybrid offspring had increased performance compared to the parental populations, as would be expected if they were genetically distinct and depauperate. Transcriptome sequencing (RNA-seq) revealed similar overall levels of genetic variation among commercial populations, but also evidence for genetic differentiation. Generally, females performed better across phenotypic traits than males. F2 hybrid offspring differed from parental populations in a highly trait-and sex specific manner. Although F2 hybrids performed better than parental populations in some traits, the results of the present study do not provide conclusive evidence that crossing of different commercial stock populations of O. majusculus improve the genetic quality and performance of this species as a biological control agent.

Prey-specific experience affects prey preference and time to kill in the soil predatory mite Gaeolaelaps aculeifer Canestrini

Biological Control, 2019

Generalist predators potentially have access to a wide array of prey, but it is little studied ho... more Generalist predators potentially have access to a wide array of prey, but it is little studied how experience with specific prey affects preference for this prey. In particular, it is unknown how experience with pest prey affects predator foraging decisions in cases where the pest is nutritious but protected by a repelling, potentially deadly defence. We investigated preference of the soil predatory mite Gaeolaelaps aculeifer Canestrini for the risky pest springtail Protaphorura fimata Gisin relative to the safe non-pest springtail Folsomia candida Willem. Egg production under foraging on live or dead individuals of either prey showed that the two species had equal nutritional quality for G. aculeifer, but indicated that live F. candida were more difficult to catch than live P. fimata. Importantly, some G. aculeifer were killed by P. fimata defence secretions, demonstrating that live P. fimata are risky prey. Preference for P. fimata was generally high when mites were given a choice between a live individual of either prey, but this preference was reduced following exposure to live individuals of P. fimata. Furthermore, fewer G. aculeifer killed a prey and time until kill was longer after experience with live P. fimata. These findings indicate that live P. fimata induced a partial aversion on G. aculeifer during exposure. Our study shows that generalist predators can reduce their preference for risky prey following exposure. This indicates that generalist predators used in biological control against risky prey are most efficient against this prey if not exposed to it prior to release.

Interactive effects of temperature and time on cold tolerance and spring predation in overwintering soil predatory mites (Gaeolaelaps aculeifer Canestrini)

Biological Control, 2019

Soil living mites have large potential as biocontrol agents against soil- dwelling pests, but lit... more Soil living mites have large potential as biocontrol agents against soil- dwelling pests, but little is known about their ecological and ecophysiological responses to cold. We investigated the interactive effects of acclimation temperature and time on cold tolerance in the laelapid mite Gaeolaelaps aculeifer Canestrini after exposure to 5, 10, 15, or 20 °C for 1, 4, or 8 days. Another group of mites were subjected to simulated winter by gradually lowering the temperature from 20 to 0.8 °C during three months, while measuring tolerance to -2 and -5 °C as well as supercooling point and melt onset temperature of body fluids at start (20 °C; “summer”), at 5 °C (“autumn”), and at 0.8 °C (“winter”). A third group was kept at constant 10 °C as a constant mild cold comparison. We found a strong interaction between exposure temperature and time on cold tolerance, with rapid cold hardening after 24 h at 5 °C but increasing cold acclimation at 10 °C. During simulated winter, tolerance to -2 °C was high after two months at 4.1 °C, but then decreased to intermediate levels after another month at 0.8 °C. The supercooling point did not change over the simulated winter, but melt onset temperature was lowered after 0.8 °C exposure. Mites preyed and reproduced readily following simulated winter, but at lower rates than if kept at constant 10 °C. Our study indicates that G. aculeifer can overwinter following release, and suggests that cold storage is advantageous before inoculative release in early spring.

Cold acclimation reduces predation rate and reproduction but increases cold- and starvation tolerance in the predatory mite Gaeolaelaps aculeifer Canestrini

Biological Control, 2017

Ectotherms respond to their thermal environments by physiological acclimation, which increases to... more Ectotherms respond to their thermal environments by physiological acclimation, which increases tolerance to thermal extremes and may increase field performance. However, acclimation often has costs, and increased performance in some traits may be associated with reduced performance in other traits due to trade-offs. We investigated effects of thermal acclimation on predation, reproduction, starvation tolerance, and locomotor activity in the predatory mite Gaeolaelaps aculeifer Canestrini at 10, 15, and 20 °C after seven days of exposure to either of the same three temperatures, following rearing at 20 °C. To test for effects of cold acclimation on cold tolerance, another set of mites were acclimated at each of the three temperatures over four days including an additional group at 5 °C, and survival was assayed following a 24 h exposure to −2 °C. Our results showed highest cold-and starvation tolerance but lowest predation and reproduction across test temperatures in mites acclimated to 10 °C. These relationships were intermediate after 15 °C acclimation and opposite after 20 °C acclimation. Locomotor activity was unaffected by acclimation temperature. Since predation and reproduction of G. aculeifer are lowered across temperatures after cold acclimation, we recommend keeping cultures at 20 °C without cold exposure until release in the field when used against heavy pest infestations. However, if using G. aculeifer to keep small infestations minimized, we recommend using cold acclimated mites that better tolerate low temperature and low prey availability. Such thermal and metabolic robustness could be advantageous if predatory control agents are introduced in agroecosystems in early spring to prevent establishment of pest populations.

Little evidence for intralocus sexual conflict over the optimal intake of nutrients for life span and reproduction in the black field cricket Teleogryllus commodus

There is often large divergence in the effects of key nutrients on life span (LS) and reproductio... more There is often large divergence in the effects of key nutrients on life span (LS) and reproduction in the sexes, yet nutrient intake is regulated in the same way in males and females given dietary choice. This suggests that the sexes are constrained from feeding to their sex-specific nutritional optima for these traits. Here, we examine the potential for intralocus sexual conflict (IASC) over optimal protein and carbohydrate intake for LS and reproduction to constrain the evolution of sex-specific nutrient regulation in the field cricket, Teleogryllus commodus. We show clear sex differences in the effects of protein and carbohydrate intake on LS and reproduction and strong positive genetic correlations between the sexes for the regulated intake of these nutrients. However, the between-sex additive genetic covariance matrix had very little effect on the predicted evolutionary response of nutrient regulation in the sexes. Thus, IASC appears unlikely to act as an evolutionary constraint on sex-specific nutrient regulation in T. commodus. This finding is supported by clear sexual dimorphism in the regulated intake of these nutrients under dietary choice. However, nutrient regulation did not coincide with the nutritional optima for LS or reproduction in either sex, suggesting that IASC is not completely resolved in T. commodus. K E Y W O R D S : Carbohydrate, Geometric Framework, intralocus sexual conflict, life span, nutrition, protein, reproduction.

Persistence of a sugar-rejecting cockroach genotype under various dietary regimes

Glucose-aversion is a heritable trait that evolved in a number of German cockroach (Blattella ger... more Glucose-aversion is a heritable trait that evolved in a number of German cockroach (Blattella germanica L.) populations in response to strong selection with glucose-containing insecticide baits. However, in the absence of glucose-containing bait, glucose-averse (GA) cockroaches have lower performance than wild-type (WT) cockroaches in several fitness-determining traits. We allocated 48 caged populations initiated with homozygous GA and WT adults to four dietary treatments consisting of either pure rodent chow, rodent chow mixed to yield a content of either 20% glucose or 20% fructose, or a treatment consisting of choice between the 20% glucose-and the 20% fructose-containing food. After 6 months we found significantly higher frequency of WT individuals in populations restricted to the 20% glucose food, and after 12 months all dietary treatments contained significantly more WT individuals than expected. In accompanying experiments, we found lower survival and longer development time of GA nymphs restricted to glucose-containing food. We furthermore found evidence for assortative mating of females with males from their own genotype, with significant differences within WT cockroaches. Our study shows experimental evidence that within heterogeneous populations, WT German cockroaches will over time prevail in abundance over GA individuals, even when glucose is not a dietary component.

The complex interplay between macronutrient intake, cuticular hydrocarbon expression and mating success in male decorated crickets

The condition dependence of male sexual traits plays a central role in sexual selection theory. R... more The condition dependence of male sexual traits plays a central role in sexual selection theory. Relatively little, however, is known about the condition dependence of chemical signals used in mate choice and their subsequent effects on male mating success. Furthermore, few studies have isolated the speciﬁc nutrients responsible for condition-dependent variation in male sexual traits. Here, we used nutritional geometry to determine the effect of protein (P) and carbohydrate (C) intake on male cuticular hydrocarbon (CHC) expression and mating success in male decorated crickets (Gryllodes sigillatus). We show that both traits are maximized at a moderate-to-high intake of nutrients in a P:C ratio of 1 : 1.5. We also show that female precopulatory mate choice exerts a complex pattern of linear and quadratic sexual selection on this condition-dependent variation in male CHC expression. Structural equation modelling revealed that although the effect of nutrient intake on mating success is mediated through condition-dependent CHC expression, it is not exclusively so, suggesting that other traits must also play an important role. Collectively, our results suggest that the complex interplay between nutrient intake, CHC expression and mating success plays an important role in the operation of sexual selection in G. sigillatus.

Effects of foraging distance on macronutrient balancing and performance in the German cockroach Blattella germainca

The German cockroach (Blattella germanica) is an excellent model omnivore for studying the effect... more The German cockroach (Blattella germanica) is an excellent model omnivore for studying the effect of foraging effort on nutrient balancing behavior and physiology, and its consequences for performance. We investigated the effect of foraging distance on individual German cockroaches by providing two foods differing in protein-to-carbohydrate ratio at opposite ends of long containers or adjacent to each other in short containers. Each food was nutritionally imbalanced, but the two foods were nutritionally complementary, allowing optimal foraging by selective feeding from both foods. We measured nutrient-specific consumption in fifth instar nymphs and newly eclosed females foraging at the two distances, hypothesizing that individuals foraging over longer distance would select more carbohydrate-biased diets to compensate for the energetic cost of locomotion. We then determined dry mass growth and lipid accumulation in the nymphs as well as mass gain and the length of basal oocytes in the adult females as an estimate of sexual maturation. Nymphs foraging over longer distance accumulated less lipid relative to total dry mass growth, but contrary to our predictions, their protein intake was higher and they accumulated more structural mass. In concordance, adult females foraging over longer distance gained more body mass and matured their oocytes faster. Our results show a positive effect of foraging distance on fitness-related parameters at two life stages, in both cases involving increased consumption of specific nutrients corresponding to requirements at the respective life stage.

Insecticide resistance and nutrition interactively shape life-history parameters in German cockroaches

Fitness-related costs of evolving insecticide resistance have been reported in a number of insect... more Fitness-related costs of evolving insecticide resistance have been reported in a number of insect species, but the interplay between evolutionary adaptation to insecticide pressure and variable environmental conditions has received little attention. We provisioned nymphs from three German cockroach (Blattella germanica L.) populations, which differed in insecticide resistance, with either nutritionally rich or poor (diluted) diet throughout their development. One population was an insecticide-susceptible laboratory strain; the other two populations originated from a field-collected indoxacarb-resistant population, which upon collection was maintained either with or without further selection with indoxacarb. We then measured development time, survival to the adult stage, adult body size, and results of a challenge with indoxacarb. Our results show that indoxacarb resistance and poor nutritional condition increased development time and lowered adult body size, with reinforcing interactions. We also found lower survival to the adult stage in the indoxacarb-selected population, which was exacerbated by poor nutrition. In addition, nutrition imparted a highly significant effect on indoxacarb susceptibility. This study exemplifies how poor nutritional condition can aggravate the life-history costs of resistance and elevate the detrimental effects of insecticide exposure, demonstrating how environmental conditions and resistance may interactively impact individual fitness and insecticide efficacy.

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

Ecology and Evolution, 2016

Sexual selection may cause dietary requirements for reproduction to diverge across the sexes and ... more 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.

Balancing of lipid, protein, and carbohydrate intake in a predatory beetle following hibernation, and consequences for lipid restoration

Carnivorous animals are known to balance their consumption of lipid and protein, and recent studi... more Carnivorous animals are known to balance their consumption of lipid and protein, and recent studies indicate that some mammalian carnivores also regulate their intake of carbohydrate. We investigated macronutrient balancing and lipid restoration following hibernation in the ground beetle Anchomenus dorsalis, hypothesizing that carbohydrates might be important energy sources upon hibernation when predator lipid stores are exhausted and prey are equally lean. We recorded the consumption of lipid, protein, and carbohydrate over nine days following hibernation, as the beetles foraged to refill their lipid stores. Each beetle was given the opportunity to regulate consumption from two semi-artificial foods differing in the proportion of two of the three macronutrients, while the third macronutrient was kept constant. When analyzing consumption of the three macronutrients on an energetic basis, it became apparent that the beetles regulated lipid and carbohydrate energy interchangeably and balanced the combined energy intake from the two macronutrients against protein intake. Restoration of lipid stores was independent of the availability of any specific macronutrient. However, the energetic consumption required to refill lipid stores was higher when a low proportion of lipids was ingested, suggesting that lipids were readily converted into lipid stores while there were energetic costs associated with converting carbohydrate and protein into stored lipids. Our experiment demonstrates that carbohydrates are consumed and regulated as a non-protein energy source by A. dorsalis despite an expectedly low occurrence of carbohydrates in their natural diet. Perhaps carbohydrates are in fact an overlooked supplementary energy source in the diet of carnivorous arthropods.

Macronutrient intake regulates sexual conflict in decorated crickets

Journal of Evolutionary Biology

Sexual conflict results in a diversity of sex-specific adaptations, including chemical additions ... more Sexual conflict results in a diversity of sex-specific adaptations, including chemical additions to ejaculates. Male decorated crickets (Gryllodes sigillatus) produce a gelatinous nuptial gift (the spermatophylax) that varies in size and free amino acid composition, which influences a female’s willingness to fully consume this gift. Complete consumption of this gift maximises sperm transfer through increased retention of the sperm-containing ampulla, but hinders post-copulatory mate choice. Here, we examine the effects of protein (P) and carbohydrate (C) intake on the weight and amino acid composition of the spermatophylax that describes its gustatory appeal to the female, as well as the ability of this gift to regulate sexual conflict via ampulla attachment time. Nutrient intake had similar effects on the expression of these traits with each maximised at a high intake of nutrients with a P:C ratio of 1:1.3. Under dietary choice, males actively regulated their nutrient intake but this regulation did not coincide with the peak of the nutritional landscape for any trait. Our results therefore demonstrate that a balanced intake of nutrients is central to regulating sexual conflict in G. sigillatus but males are constrained from reaching the optima needed to bias the outcome of this conflict in their favour.

The geometry of nutrient space-based life-history trade-offs: sex-specific effects of macronutrient intake on the trade-off between encapsulation ability and reproductive effort in decorated crickets

American Naturalist, 2018

Life-history theory assumes that traits compete for limited resources, resulting in trade-offs. T... more Life-history theory assumes that traits compete for limited resources, resulting in trade-offs. The most commonly manipulated resource in empirical studies is the quantity or quality of diet. Recent studies using the geometric framework for nutrition, however, suggest that trade-offs are often regulated by the intake of specific nutrients, but a formal approach to identify and quantify the strength of such trade-offs is lacking. We posit that trade-offs occur whenever life-history traits are maximized in different regions of nutrient space, as evidenced by nonoverlapping 95% confidence regions of the global maximum for each trait and large angles (v) between linear nutritional vectors and Euclidean distances (d) between global maxima. We then examined the effects of protein and carbohydrate intake on the tradeoff between reproduction and aspects of immune function in male and female Gryllodes sigillatus. Female encapsulation ability and egg production increased with the intake of both nutrients, whereas male encapsulation ability increased with protein intake but calling effort increased with carbohydrate intake. The trade-offs between traits was therefore larger in males than in females, as demonstrated by significant negative correlations between the traits in males, nonoverlapping 95% confidence regions, and larger estimates of v and d. Under dietary choice, the sexes had similar regulated intakes, but neither optimally regulated nutrient intake for maximal trait expression. We highlight the fact that greater consideration of specific nutrient intake is needed when examining nutrient space-based trade-offs.

Intralocus sexual conflict over optimal nutrient intake and the evolution of sex differences in life span and reproduction

Functional Ecology, 2022

1. Despite widespread variation in life span across species, three clear patterns exist: sex diff... more 1. Despite widespread variation in life span across species, three clear patterns exist: sex differences in life span are ubiquitous, life span is commonly traded against reproduction, and nutrition has a major influence on these traits and how they trade-off. One process that potentially unites these patterns is intralocus sexual conflict over the optimal intake of nutrients for life span and reproduction. If nutrient intake has sex-specific effects on life span and reproduction but nutrient choice is genetically linked across the sexes, intralocus sexual conflict will occur and may prevent one or both sexes from feeding to their nutritional optima.
2. Here we determine the potential for this process to operate in the decorated cricket Gryllodes sigillatus. Using the Geometric Framework for Nutrition, we restrict male and female crickets to diets varying in the ratio of protein to carbohydrates and total nutrient content to quantify the effects on life span and daily reproductive effort in the sexes. We then use inbred lines to estimate the quantitative genetic basis of nutrient choice in males and females. We combine the nutrient effects and genetic estimates to predict the magnitude of evolutionary constraint for these traits in each sex. Finally, we present male and female crickets with a much broader range of diet pairs to determine how the sexes actively regulate their intake of nutrients.
3. We show that protein and carbohydrate intake have contrasting effects on life span and reproduction in the sexes and that there are strong positive intersexual genetic correlations for the intake of these nutrients under dietary choice. This is predicted to accelerate the evolutionary response of nutrient intake in males but constrain it in females, suggesting they are losing the conflict. Supporting this view, males and females regulate nutrient intake to a common nutrient ratio that was not perfectly optimal for life span or reproduction in either sex, especially in females.
4. Our findings show that intralocus sexual conflict over the optimal intake of nutrients is likely to be an important process generating sex differences in life span and reproduction and may help explain why females age faster and live shorter than males in G. sigillatus.

Rapid evolutionary adaptation to diet composition in the black soldier fly (Hermetia illucens)

Insects, 2023

Genetic adaptation of Hermetia illucens (BSF) to suboptimal single sourced waste streams can open... more Genetic adaptation of Hermetia illucens (BSF) to suboptimal single sourced waste streams can open new perspectives for insect production. Here, four BSF lines were maintained on a single sourced, low-quality wheat bran diet (WB) or on a high-quality chicken feed diet (CF) for 13 generations. We continuously evaluated presumed evolutionary responses in several performance traits to rearing on the two diets. Subsequently, we tested responses to interchanged diets, i.e., of larvae that had been reared on low-quality feed and tested on high-quality feed and vice versa to evaluate costs associated with adaptation to different diets. BSF were found to experience rapid adaptation to the diet composition. While performances on the WB diet were always inferior to the CF diet, the adaptive responses were stronger to the former diet. This stronger response was likely due to stronger selection pressure experienced by BSF fed on the low-quality single sourced diet. The interchanged diet experiment found no costs associated with diet adaptation, but revealed cross generational gain associated with the parental CF diet treatment. Our results revealed that BSF can rapidly respond adaptively to diet, although the mechanisms are yet to be determined. This has potential to be utilized in commercial insect breeding to produce lines tailored to specific diets.

Pilot scale production of Hermetia illucens (L.) larvae and frass using former foodstuffs

Cleaner Engineering and Technology, 2022

The food and feed sector requires new sustainable sources of protein and innovative solutions for... more The food and feed sector requires new sustainable sources of protein and innovative solutions for upcycling of food waste (former foodstuffs), which today is downcycled into energy or even wasted. This study aimed at evaluating the use of former foodstuff waste streams as feed substrate for Hermetia illucens (L.) larvae (black soldier fly larvae, BSFL) under long-term and semi-industrial conditions. Different foodstuff-based mixtures and different stocking BSFL densities were used during 20 batches, and quality and safety assessments were performed on the main outputs, namely BSFL production performance, frass impurities, larval and frass nutrient profiles and heavy metal content. About 1400 kg of former foodstuffs (fresh weight) were used to produce 239 kg BSFL and 230 kg frass. The production of BSFL reared on former foodstuffs was highly efficient, with feed conversion rates (FCR) ranging between 2.3 and 5.5 (dry matter basis). The optimization experiment revealed that former foodstuffs-based mixture and high larval density (10 larvae/cm 2) lead to highly efficient (FCR: 2.6) and heavy metal-free production of BSFL and frass. The quality of the derived BSFL meal was high in terms of protein and amino acids. Furthermore, the quality of the technical frass was high in terms of N, P, and K levels and minimal packaging material residuals (<2.65%). This investigation suggests that nutrients in former foodstuffs can be successfully and safely recycled in production of BSFL.

Genotype-by-sex-by-diet interactions for nutritional preference, dietary consumption, and lipid deposition in a field cricket

Heredity, 2018

Changes in feeding behaviour, especially the overconsumption of calories, has led to a rise in th... more Changes in feeding behaviour, especially the overconsumption of calories, has led to a rise in the rates of obesity, diabetes, and other associated disorders in humans and a range of animals inhabiting human-influenced environments. However, understanding the relative contribution of genes, the nutritional environment, and their interaction to dietary intake and lipid deposition in the sexes still remains a major challenge. By combining nutritional geometry with quantitative genetics, we determined the effect of genes, the nutritional environment, and their interaction on the total nutritional preference (TP), total diet eaten (TE), and lipid mass (LM) of male and female black field crickets (Teleogryllus commodus) fed one of four diet pairs (DPs) differing in the ratio of protein to carbohydrate and total nutritional content. We found abundant additive genetic variance for TP, TE, and LM in both sexes and across all four DPs, with significant genetic correlations between TE and TP ...

Geometric Stoichiometry: unifying concepts of animal nutrition to understand how protein-rich diets can be "too much of a good thing"

Frontiers in Ecology and Evolution, 2020

Understanding the factors that control the growth of heterotrophic organisms is central to predic... more Understanding the factors that control the growth of heterotrophic organisms is central to predicting food web interactions and biogeochemical cycling within ecosystems. We present a new framework, Geometric Stoichiometry (GS), that unifies the disciplines of Nutritional Geometry (NG) and Ecological Stoichiometry (ES) by extending the equations of ES to incorporate core NG concepts, including macromolecules as currencies and the ability of animals to select foods that balance deficits and excesses of nutrients. The resulting model is used to investigate regulation of consumer growth by dietary protein:carbohydrate ratio. Growth on protein-poor diets is limited by nitrogen. Likewise, we show that growth is also diminished on protein-rich diets and that this can be mechanistically explained by means of a metabolic penalty that arises when animals use protein for energy generation. These penalties, which are incurred when dealing with the costs of producing and excreting toxic nitrogenous waste, have not hitherto been represented in standard ES theory. In order to incorporate GS within ecosystem and biogeochemical models, a new generation of integrated theoretical and experimental studies based on unified concepts of NG and ES is needed, including measurements of food selection, biomass, growth and associated physiology, and involving metabolic penalties.

Prey-specific impact of cold pre-exposure on kill rate and reproduction

Journal of Animal Ecology, 2019

1. Temperature influences biological processes of ectotherms including ecological interactions, b... more 1. Temperature influences biological processes of ectotherms including ecological interactions, but interaction strengths may depend on species-specific traits. Furthermore, ectotherms acclimate to prevailing thermal conditions by adjusting physiological parameters, which often implies costs to other fitness-related parameters. Both predators and prey may therefore pay thermal acclimation costs following exposure to suboptimal temperatures. However, these costs may be asymmetrical between predator and prey, and between the predator and different species of concurrent prey. 2. We investigated whether thermal pre-exposure affected subsequent kill rate and predator fitness when foraging on prey that differ in ease of capture, and whether changes were primarily caused by predator or by prey pre-exposure effects. Specifically, we were interested in whether there were interactions between predator pre-exposed temperature and specific prey. 3. Using the mesostigmatid mite Gaeolaelaps aculeifer as a generalist predator and the collembolans Folsomia candida and Protaphorura fimata as prey, we measured the impact of present temperature, predator pre-exposure temperature, prey pre-exposure temperature (all 10 or 20°C), prey species, and all interactions on prey numbers killed, predator eggs produced, and exploitation of killed prey in a full factorial design. 4. Mites killed P. fimata in equal numbers independent of the presence of F. candida, but killed F. candida when P. fimata was absent. Mite kill rate and reproduction were significantly affected by mite pre-exposure temperature and test temperature , but not by prey pre-exposure temperature. Significantly more of the slower prey was killed than of the quicker prey. Importantly, we found significant syner-gistic negative interaction effects between predator cold pre-exposure and hunting prey of higher agility on predator kill rate and reproduction. 5. Our findings show that the negative effects of cold and cold pre-exposure on kill rate and reproduction may be more severe when predators forage on quick prey. The study implies that predator cold exposure has consequences for specific prey survival following cold due to altered predation pressures, which in nature should influence the specific prey population dynamics and apparent competition

Increased lipid accumulation but not reduced metabolism explains improved starvation tolerance in cold-acclimated arthropod predators

Science of Nature, 2018

Predatory arthropods are used for biological control in greenhouses, but there is increasing inte... more Predatory arthropods are used for biological control in greenhouses, but there is increasing interest to extend their use to the outdoor environment where temperatures are typically lower. Acclimation at low temperature increases the ability of ectotherms to cope with subsequent more extreme cold, but may involve costs or benefits to other performance traits. A recent study in mesostigmatid mites (Gaeolaelaps aculeifer) showed that starvation tolerance was improved following a period of cold exposure. However, the physiological mechanisms that underlie improved starvation tolerance following cold exposure were not investigated. To examine whether cold acclimation would also improve starvation tolerance in an insect, we repeated the starvation study in another arthropod predator, the pirate bug Orius majusculus, as well as in G. aculeifer. Before tests, the two species were acclimated at 10, 15, or 20°C for 7 (G. aculeifer) or 16 (O. majusculus) days. We then analyzed the effects of thermal exposure on body composition, consumption, and basal metabolic rate in both species. Our results confirmed that exposure to low temperature improves starvation tolerance in these arthropod predators. Body composition analyses revealed that both species had accumulated larger lipid stores during exposure to colder temperature, which at least in part can explain the larger starvation tolerance following cold exposure. In contrast, consumption and basal metabolic rate were not changed by thermal acclimation. Our study indicates that predatory arthropods exposed to cold increase their physiological robustness and ability to endure environmental challenges, including low temperature and low prey availability.

Are commercial stocks of biological control agents genetically depauperate? -A case study on the pirate bug Orius majusculus Reuter

Biological Control, 2018

Predatory arthropods are increasingly used in biological control of insect pests. For this purpos... more Predatory arthropods are increasingly used in biological control of insect pests. For this purpose, control agents are produced commercially in large quantities for release in crops. The production stocks, however, may have undergone numerous population bottlenecks and may have been exposed to artificial selection pressures in the production facilities. Accordingly, commercial populations may be experiencing loss of genetic variation through inbreeding and genetic drift, which may reduce fitness and biocontrol efficiency. In the present study we investigated whether populations of the pirate bug Orius majusculus (Reuter) purchased from three European companies differed in a range of performance traits including predation rate, starvation tolerance, body size, locomotor activity, and heat tolerance. Furthermore, we crossed all populations pairwise and tested whether outcrossed F2 hybrid offspring had increased performance compared to the parental populations, as would be expected if they were genetically distinct and depauperate. Transcriptome sequencing (RNA-seq) revealed similar overall levels of genetic variation among commercial populations, but also evidence for genetic differentiation. Generally, females performed better across phenotypic traits than males. F2 hybrid offspring differed from parental populations in a highly trait-and sex specific manner. Although F2 hybrids performed better than parental populations in some traits, the results of the present study do not provide conclusive evidence that crossing of different commercial stock populations of O. majusculus improve the genetic quality and performance of this species as a biological control agent.

Prey-specific experience affects prey preference and time to kill in the soil predatory mite Gaeolaelaps aculeifer Canestrini

Biological Control, 2019

Generalist predators potentially have access to a wide array of prey, but it is little studied ho... more Generalist predators potentially have access to a wide array of prey, but it is little studied how experience with specific prey affects preference for this prey. In particular, it is unknown how experience with pest prey affects predator foraging decisions in cases where the pest is nutritious but protected by a repelling, potentially deadly defence. We investigated preference of the soil predatory mite Gaeolaelaps aculeifer Canestrini for the risky pest springtail Protaphorura fimata Gisin relative to the safe non-pest springtail Folsomia candida Willem. Egg production under foraging on live or dead individuals of either prey showed that the two species had equal nutritional quality for G. aculeifer, but indicated that live F. candida were more difficult to catch than live P. fimata. Importantly, some G. aculeifer were killed by P. fimata defence secretions, demonstrating that live P. fimata are risky prey. Preference for P. fimata was generally high when mites were given a choice between a live individual of either prey, but this preference was reduced following exposure to live individuals of P. fimata. Furthermore, fewer G. aculeifer killed a prey and time until kill was longer after experience with live P. fimata. These findings indicate that live P. fimata induced a partial aversion on G. aculeifer during exposure. Our study shows that generalist predators can reduce their preference for risky prey following exposure. This indicates that generalist predators used in biological control against risky prey are most efficient against this prey if not exposed to it prior to release.

Interactive effects of temperature and time on cold tolerance and spring predation in overwintering soil predatory mites (Gaeolaelaps aculeifer Canestrini)

Biological Control, 2019

Soil living mites have large potential as biocontrol agents against soil- dwelling pests, but lit... more Soil living mites have large potential as biocontrol agents against soil- dwelling pests, but little is known about their ecological and ecophysiological responses to cold. We investigated the interactive effects of acclimation temperature and time on cold tolerance in the laelapid mite Gaeolaelaps aculeifer Canestrini after exposure to 5, 10, 15, or 20 °C for 1, 4, or 8 days. Another group of mites were subjected to simulated winter by gradually lowering the temperature from 20 to 0.8 °C during three months, while measuring tolerance to -2 and -5 °C as well as supercooling point and melt onset temperature of body fluids at start (20 °C; “summer”), at 5 °C (“autumn”), and at 0.8 °C (“winter”). A third group was kept at constant 10 °C as a constant mild cold comparison. We found a strong interaction between exposure temperature and time on cold tolerance, with rapid cold hardening after 24 h at 5 °C but increasing cold acclimation at 10 °C. During simulated winter, tolerance to -2 °C was high after two months at 4.1 °C, but then decreased to intermediate levels after another month at 0.8 °C. The supercooling point did not change over the simulated winter, but melt onset temperature was lowered after 0.8 °C exposure. Mites preyed and reproduced readily following simulated winter, but at lower rates than if kept at constant 10 °C. Our study indicates that G. aculeifer can overwinter following release, and suggests that cold storage is advantageous before inoculative release in early spring.

Cold acclimation reduces predation rate and reproduction but increases cold- and starvation tolerance in the predatory mite Gaeolaelaps aculeifer Canestrini

Biological Control, 2017

Ectotherms respond to their thermal environments by physiological acclimation, which increases to... more Ectotherms respond to their thermal environments by physiological acclimation, which increases tolerance to thermal extremes and may increase field performance. However, acclimation often has costs, and increased performance in some traits may be associated with reduced performance in other traits due to trade-offs. We investigated effects of thermal acclimation on predation, reproduction, starvation tolerance, and locomotor activity in the predatory mite Gaeolaelaps aculeifer Canestrini at 10, 15, and 20 °C after seven days of exposure to either of the same three temperatures, following rearing at 20 °C. To test for effects of cold acclimation on cold tolerance, another set of mites were acclimated at each of the three temperatures over four days including an additional group at 5 °C, and survival was assayed following a 24 h exposure to −2 °C. Our results showed highest cold-and starvation tolerance but lowest predation and reproduction across test temperatures in mites acclimated to 10 °C. These relationships were intermediate after 15 °C acclimation and opposite after 20 °C acclimation. Locomotor activity was unaffected by acclimation temperature. Since predation and reproduction of G. aculeifer are lowered across temperatures after cold acclimation, we recommend keeping cultures at 20 °C without cold exposure until release in the field when used against heavy pest infestations. However, if using G. aculeifer to keep small infestations minimized, we recommend using cold acclimated mites that better tolerate low temperature and low prey availability. Such thermal and metabolic robustness could be advantageous if predatory control agents are introduced in agroecosystems in early spring to prevent establishment of pest populations.

Little evidence for intralocus sexual conflict over the optimal intake of nutrients for life span and reproduction in the black field cricket Teleogryllus commodus

There is often large divergence in the effects of key nutrients on life span (LS) and reproductio... more There is often large divergence in the effects of key nutrients on life span (LS) and reproduction in the sexes, yet nutrient intake is regulated in the same way in males and females given dietary choice. This suggests that the sexes are constrained from feeding to their sex-specific nutritional optima for these traits. Here, we examine the potential for intralocus sexual conflict (IASC) over optimal protein and carbohydrate intake for LS and reproduction to constrain the evolution of sex-specific nutrient regulation in the field cricket, Teleogryllus commodus. We show clear sex differences in the effects of protein and carbohydrate intake on LS and reproduction and strong positive genetic correlations between the sexes for the regulated intake of these nutrients. However, the between-sex additive genetic covariance matrix had very little effect on the predicted evolutionary response of nutrient regulation in the sexes. Thus, IASC appears unlikely to act as an evolutionary constraint on sex-specific nutrient regulation in T. commodus. This finding is supported by clear sexual dimorphism in the regulated intake of these nutrients under dietary choice. However, nutrient regulation did not coincide with the nutritional optima for LS or reproduction in either sex, suggesting that IASC is not completely resolved in T. commodus. K E Y W O R D S : Carbohydrate, Geometric Framework, intralocus sexual conflict, life span, nutrition, protein, reproduction.

Persistence of a sugar-rejecting cockroach genotype under various dietary regimes

Glucose-aversion is a heritable trait that evolved in a number of German cockroach (Blattella ger... more Glucose-aversion is a heritable trait that evolved in a number of German cockroach (Blattella germanica L.) populations in response to strong selection with glucose-containing insecticide baits. However, in the absence of glucose-containing bait, glucose-averse (GA) cockroaches have lower performance than wild-type (WT) cockroaches in several fitness-determining traits. We allocated 48 caged populations initiated with homozygous GA and WT adults to four dietary treatments consisting of either pure rodent chow, rodent chow mixed to yield a content of either 20% glucose or 20% fructose, or a treatment consisting of choice between the 20% glucose-and the 20% fructose-containing food. After 6 months we found significantly higher frequency of WT individuals in populations restricted to the 20% glucose food, and after 12 months all dietary treatments contained significantly more WT individuals than expected. In accompanying experiments, we found lower survival and longer development time of GA nymphs restricted to glucose-containing food. We furthermore found evidence for assortative mating of females with males from their own genotype, with significant differences within WT cockroaches. Our study shows experimental evidence that within heterogeneous populations, WT German cockroaches will over time prevail in abundance over GA individuals, even when glucose is not a dietary component.

The complex interplay between macronutrient intake, cuticular hydrocarbon expression and mating success in male decorated crickets

The condition dependence of male sexual traits plays a central role in sexual selection theory. R... more The condition dependence of male sexual traits plays a central role in sexual selection theory. Relatively little, however, is known about the condition dependence of chemical signals used in mate choice and their subsequent effects on male mating success. Furthermore, few studies have isolated the speciﬁc nutrients responsible for condition-dependent variation in male sexual traits. Here, we used nutritional geometry to determine the effect of protein (P) and carbohydrate (C) intake on male cuticular hydrocarbon (CHC) expression and mating success in male decorated crickets (Gryllodes sigillatus). We show that both traits are maximized at a moderate-to-high intake of nutrients in a P:C ratio of 1 : 1.5. We also show that female precopulatory mate choice exerts a complex pattern of linear and quadratic sexual selection on this condition-dependent variation in male CHC expression. Structural equation modelling revealed that although the effect of nutrient intake on mating success is mediated through condition-dependent CHC expression, it is not exclusively so, suggesting that other traits must also play an important role. Collectively, our results suggest that the complex interplay between nutrient intake, CHC expression and mating success plays an important role in the operation of sexual selection in G. sigillatus.

Effects of foraging distance on macronutrient balancing and performance in the German cockroach Blattella germainca

The German cockroach (Blattella germanica) is an excellent model omnivore for studying the effect... more The German cockroach (Blattella germanica) is an excellent model omnivore for studying the effect of foraging effort on nutrient balancing behavior and physiology, and its consequences for performance. We investigated the effect of foraging distance on individual German cockroaches by providing two foods differing in protein-to-carbohydrate ratio at opposite ends of long containers or adjacent to each other in short containers. Each food was nutritionally imbalanced, but the two foods were nutritionally complementary, allowing optimal foraging by selective feeding from both foods. We measured nutrient-specific consumption in fifth instar nymphs and newly eclosed females foraging at the two distances, hypothesizing that individuals foraging over longer distance would select more carbohydrate-biased diets to compensate for the energetic cost of locomotion. We then determined dry mass growth and lipid accumulation in the nymphs as well as mass gain and the length of basal oocytes in the adult females as an estimate of sexual maturation. Nymphs foraging over longer distance accumulated less lipid relative to total dry mass growth, but contrary to our predictions, their protein intake was higher and they accumulated more structural mass. In concordance, adult females foraging over longer distance gained more body mass and matured their oocytes faster. Our results show a positive effect of foraging distance on fitness-related parameters at two life stages, in both cases involving increased consumption of specific nutrients corresponding to requirements at the respective life stage.

Insecticide resistance and nutrition interactively shape life-history parameters in German cockroaches

Fitness-related costs of evolving insecticide resistance have been reported in a number of insect... more Fitness-related costs of evolving insecticide resistance have been reported in a number of insect species, but the interplay between evolutionary adaptation to insecticide pressure and variable environmental conditions has received little attention. We provisioned nymphs from three German cockroach (Blattella germanica L.) populations, which differed in insecticide resistance, with either nutritionally rich or poor (diluted) diet throughout their development. One population was an insecticide-susceptible laboratory strain; the other two populations originated from a field-collected indoxacarb-resistant population, which upon collection was maintained either with or without further selection with indoxacarb. We then measured development time, survival to the adult stage, adult body size, and results of a challenge with indoxacarb. Our results show that indoxacarb resistance and poor nutritional condition increased development time and lowered adult body size, with reinforcing interactions. We also found lower survival to the adult stage in the indoxacarb-selected population, which was exacerbated by poor nutrition. In addition, nutrition imparted a highly significant effect on indoxacarb susceptibility. This study exemplifies how poor nutritional condition can aggravate the life-history costs of resistance and elevate the detrimental effects of insecticide exposure, demonstrating how environmental conditions and resistance may interactively impact individual fitness and insecticide efficacy.

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

Ecology and Evolution, 2016

Sexual selection may cause dietary requirements for reproduction to diverge across the sexes and ... more 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.

Balancing of lipid, protein, and carbohydrate intake in a predatory beetle following hibernation, and consequences for lipid restoration

Carnivorous animals are known to balance their consumption of lipid and protein, and recent studi... more Carnivorous animals are known to balance their consumption of lipid and protein, and recent studies indicate that some mammalian carnivores also regulate their intake of carbohydrate. We investigated macronutrient balancing and lipid restoration following hibernation in the ground beetle Anchomenus dorsalis, hypothesizing that carbohydrates might be important energy sources upon hibernation when predator lipid stores are exhausted and prey are equally lean. We recorded the consumption of lipid, protein, and carbohydrate over nine days following hibernation, as the beetles foraged to refill their lipid stores. Each beetle was given the opportunity to regulate consumption from two semi-artificial foods differing in the proportion of two of the three macronutrients, while the third macronutrient was kept constant. When analyzing consumption of the three macronutrients on an energetic basis, it became apparent that the beetles regulated lipid and carbohydrate energy interchangeably and balanced the combined energy intake from the two macronutrients against protein intake. Restoration of lipid stores was independent of the availability of any specific macronutrient. However, the energetic consumption required to refill lipid stores was higher when a low proportion of lipids was ingested, suggesting that lipids were readily converted into lipid stores while there were energetic costs associated with converting carbohydrate and protein into stored lipids. Our experiment demonstrates that carbohydrates are consumed and regulated as a non-protein energy source by A. dorsalis despite an expectedly low occurrence of carbohydrates in their natural diet. Perhaps carbohydrates are in fact an overlooked supplementary energy source in the diet of carnivorous arthropods.

Macronutrient intake regulates sexual conflict in decorated crickets

Journal of Evolutionary Biology

Sexual conflict results in a diversity of sex-specific adaptations, including chemical additions ... more Sexual conflict results in a diversity of sex-specific adaptations, including chemical additions to ejaculates. Male decorated crickets (Gryllodes sigillatus) produce a gelatinous nuptial gift (the spermatophylax) that varies in size and free amino acid composition, which influences a female’s willingness to fully consume this gift. Complete consumption of this gift maximises sperm transfer through increased retention of the sperm-containing ampulla, but hinders post-copulatory mate choice. Here, we examine the effects of protein (P) and carbohydrate (C) intake on the weight and amino acid composition of the spermatophylax that describes its gustatory appeal to the female, as well as the ability of this gift to regulate sexual conflict via ampulla attachment time. Nutrient intake had similar effects on the expression of these traits with each maximised at a high intake of nutrients with a P:C ratio of 1:1.3. Under dietary choice, males actively regulated their nutrient intake but this regulation did not coincide with the peak of the nutritional landscape for any trait. Our results therefore demonstrate that a balanced intake of nutrients is central to regulating sexual conflict in G. sigillatus but males are constrained from reaching the optima needed to bias the outcome of this conflict in their favour.