Age-dependent variation in the terminal investment threshold in male crickets

This is the author accepted manuscript, made available online by the publisher 1st February 2018. Final version to be available from the publisher via the DOI in this record.The terminal investment hypothesis proposes that decreased expectation of future reproduction (e.g., arising from a threat to survival) should precipitate increased investment in current reproduction. The level at which a cue of decreased survival is sufficient to trigger terminal investment (i.e., the terminal investment threshold) may vary according to other factors that influence expectation for future reproduction. We test whether the terminal investment threshold varies with age in male crickets, using heat-killed bacteria to simulate an immune-inducing infection. We measured calling effort (a behavior essential for mating) and hemolymph antimicrobial activity in young and old males across a gradient of increasing infection cue intensity. There was a significant interaction between the infection cue and age in their effect on calling effort, confirming the existence of a dynamic terminal investment threshold: young males reduced effort at all infection levels, whereas old males increased effort at the highest levels relative to naïve individuals. A lack of a corresponding decrease in antibacterial activity suggests that altered reproductive effort is not traded against investment in this component of immunity. Collectively, these results support the existence of a dynamic terminal investment threshold, perhaps accounting for some of the conflicting evidence in support of terminal investment. This article is protected by copyright. All rights reserved.This research was funded, in part, by a grant from the National Science Foundation IOS 16-54028 (SKS, BMS, and JH), grants from the Beta Lambda Chapter of the Phi Sigma Biological Honor Society, Graduate Student Association of Illinois State University, Animal Behavior Society, and Orthopterists‘ Society to KRD, and an Illinois State University Summer Faculty Fellowship and Faculty Research Award to SKS

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

This is the final version of the article. Available from University of Chicago Press via the DOI in this record.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 (θ) between linear nutritional vectors and Euclidean distances (d) between global maxima. We then examined the effects of protein and carbohydrate intake on the trade-off 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 θ 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.S.K.S. was funded by the National Science Foundation (NSF; IOS-1118160 and IOS-1654028). C.M.H. was funded by a Leverhulme Early Career Fellowship. J.H. was funded by a Royal Society Fellowship (UF120087) and Equipment Grant (RG090854) and by the Natural Environment Research Council (NERC; NE/G00949X/1). J.R. was funded by a NERC studentship (awarded to J.H.). E.D.C. was partially funded by NSF grant CMII 1634878

What's in the Gift? Towards a Molecular Dissection of Nuptial Feeding in a Cricket.

Journal ArticleResearch Support, Non-U.S. Gov'tResearch Support, U.S. Gov't, Non-P.H.S.Nuptial gifts produced by males and transferred to females during copulation are common in insects. Yet, their precise composition and subsequent physiological effects on the female recipient remain unresolved. Male decorated crickets Gryllodes sigillatus transfer a spermatophore to the female during copulation that is composed of an edible gift, the spermatophylax, and the ampulla that contains the ejaculate. After transfer of the spermatophore, the female detaches the spermatophylax and starts to eat it while sperm from the ampulla are evacuated into the female reproductive tract. When the female has finished consuming the spermatophylax, she detaches the ampulla and terminates sperm transfer. Hence, one simple function of the spermatophylax is to ensure complete sperm transfer by distracting the female from prematurely removing the ampulla. However, the majority of orally active components of the spermatophylax itself and their subsequent effects on female behavior have not been identified. Here, we report the first analysis of the proteome of the G. sigillatus spermatophylax and the transcriptome of the male accessory glands that make these proteins. The accessory gland transcriptome was assembled into 17,691 transcripts whilst about 30 proteins were detected within the mature spermatophylax itself. Of these 30 proteins, 18 were encoded by accessory gland encoded messages. Most spermatophylax proteins show no similarity to proteins with known biological functions and are therefore largely novel. A spermatophylax protein shows similarity to protease inhibitors suggesting that it may protect the biologically active components from digestion within the gut of the female recipient. Another protein shares similarity with previously characterized insect polypeptide growth factors suggesting that it may play a role in altering female reproductive physiology concurrent with fertilization. Characterization of the spermatophylax proteome provides the first step in identifying the genes encoding these proteins in males and in understanding their biological functions in the female recipient.Max Planck GesellschaftNational Science FoundationBBSRRoyal Societ

Large Numbers of Matings Give Female Field Crickets a Direct Benefit but not a Genetic Benefit

Female crickets can potentially gain both direct and indirect benefits from mating multiple times with different males. Most studies have only examined the effects of small numbers of matings, although female crickets are capable of mating many times. The goal of this paper is to examine the direct and indirect benefits of mating large numbers of times for female reproductive success. In a previous experiment, female Gryllus vocalis were found to gain diminishing direct benefits from mating large numbers of times. In this study I attempt to determine whether mating large numbers of times yields similar diminishing returns on female indirect benefits. Virgin female Gryllus vocalis crickets were assigned to mate five, ten or 15 times with either the same or different males. Females that mated more times gained direct benefits in terms of laying more eggs and more fertilized eggs. Females that mated with different males rather than mating repeatedly with the same male did not have higher offspring hatching success, a result that is contrary to other published results comparing female reproductive success with repeated versus different partners. These results suggest that females that mate large numbers of times fail to gain additional genetic benefits from doing so

Genetic and environmental variation in condition, cutaneous immunity, and haematocrit in house wrens

This is the final version of the article. Available from BioMed Central via the DOI in this record.BACKGROUND: Life-history studies of wild bird populations often focus on the relationship between an individual's condition and its capacity to mount an immune response, as measured by a commonly-employed assay of cutaneous immunity, the PHA skin test. In addition, haematocrit, the packed cell volume in relation to total blood volume, is often measured as an indicator of physiological performance. A multi-year study of a wild population of house wrens has recently revealed that those exhibiting the highest condition and strongest PHA responses as nestlings are most likely to be recruited to the breeding population and to breed through two years of age; in contrast, intermediate haematocrit values result in the highest recruitment to the population. Selection theory would predict, therefore, that most of the underlying genetic variation in these traits should be exhausted resulting in low heritability, although such traits may also exhibit low heritability because of increased residual variance. Here, we examine the genetic and environmental variation in condition, cutaneous immunity, and haematocrit using an animal model based on a pedigree of approximately 2,800 house wrens. RESULTS: Environmental effects played a paramount role in shaping the expression of the fitness-related traits measured in this wild population, but two of them, condition and haematocrit, retained significant heritable variation. Condition was also positively correlated with both the PHA response and haematocrit, but in the absence of any significant genetic correlations, it appears that this covariance arises through parallel effects of the environment acting on this suite of traits. CONCLUSIONS: The maintenance of genetic variation in different measures of condition appears to be a pervasive feature of wild bird populations, in contradiction of conventional selection theory. A major challenge in future studies will be to explain how such variation persists in the face of the directional selection acting on condition in house wrens and other species.We thank the 2004–2006 Wren Crews for field assistance and the ParkLands Foundation (Merwin Preserve) and the Sears and Butler families for the use of their properties. Financial support was provided by NSF grants GK12-0086354, IBN-0316580, IOS-0718140 and IOS-1118160; NIH grant R15HD076308-01; a visiting professorship from the Leverhulme Trust (SKS); the School of Biological Sciences, Illinois State University; a BBSRC David Phillips Research Fellowship (AJW); and student-research grants from the Beta Lambda Chapter of the Phi Sigma Biological Sciences Honor Society (AMF)

Female Burying Beetles Benefit from Male Desertion: Sexual Conflict and Counter-Adaptation over Parental Investment

Sexual conflict drives the coevolution of sexually antagonistic traits, such that an adaptation in one sex selects an opposing coevolutionary response from the other. Although many adaptations and counteradaptations have been identified in sexual conflict over mating interactions, few are known for sexual conflict over parental investment. Here we investigate a possible coevolutionary sequence triggered by mate desertion in the burying beetle Nicrophorus vespilloides, where males commonly leave before their offspring reach independence. Rather than suffer fitness costs as a consequence, our data suggest that females rely on the male's absence to recoup some of the costs of larval care, presumably because they are then free to feed themselves on the carcass employed for breeding. Consequently, forcing males to stay until the larvae disperse reduces components of female fitness to a greater extent than caring for young singlehandedly. Therefore we suggest that females may have co-evolved to anticipate desertion by their partners so that they now benefit from the male's absence

Cost of Mating and Insemination Capacity of a Genetically Modified Mosquito Aedes aegypti OX513A Compared to Its Wild Type Counterpart

The idea of implementing genetics-based insect control strategies modelled on the traditional SIT is becoming increasingly popular. In this paper we compare a genetically modified line of Aedes aegypti carrying a tetracycline repressible, lethal positive feedback system (OX513A) with its wild type counterpart with respect to their insemination capacities and the cost of courtship and mating. Genetically modified males inseminated just over half as many females as the wild type males during their lifetime. Providing days of rest from mating had no significant effect on the total number of females inseminated by males of either line, but it did increase their longevity. Producing sperm had a low cost in terms of energy investment; the cost of transferring this sperm to a receptive female was much higher. Continued mating attempts with refractory females suggest that males could not identify refractory females before investing substantial energy in courtship. Although over a lifetime OX513A males inseminated fewer females, the number of females inseminated over the first three days, was similar between males of the two lines, suggesting that the identified cost of RIDL may have little impact on the outcome of SIT-based control programmes with frequent releases of the genetically modified males

Female Sexual Polymorphism and Fecundity Consequences of Male Mating Harassment in the Wild

Genetic and phenotypic variation in female response towards male mating attempts has been found in several laboratory studies, demonstrating sexually antagonistic co-evolution driven by mating costs on female fitness. Theoretical models suggest that the type and degree of genetic variation in female resistance could affect the evolutionary outcome of sexually antagonistic mating interactions, resulting in either rapid development of reproductive isolation and speciation or genetic clustering and female sexual polymorphisms. However, evidence for genetic variation of this kind in natural populations of non-model organisms is very limited. Likewise, we lack knowledge on female fecundity-consequences of matings and the degree of male mating harassment in natural settings. Here we present such data from natural populations of a colour polymorphic damselfly. Using a novel experimental technique of colour dusting males in the field, we show that heritable female colour morphs differ in their propensity to accept male mating attempts. These morphs also differ in their degree of resistance towards male mating attempts, the number of realized matings and in their fecundity-tolerance to matings and mating attempts. These results show that there may be genetic variation in both resistance and tolerance to male mating attempts (fitness consequences of matings) in natural populations, similar to the situation in plant-pathogen resistance systems. Male mating harassment could promote the maintenance of a sexual mating polymorphism in females, one of few empirical examples of sympatric genetic clusters maintained by sexual conflict

Sperm transfer or spermatangia removal: postcopulatory behaviour of picking up spermatangium by female Japanese pygmy squid

In the Japanese pygmy squid Idiosepius paradoxus, females often pick up the spermatangium using their mouth (buccal mass) after copulation. To examine whether the female I. paradoxus directly transfers sperm into the seminal receptacle via this picking behaviour, or removes the spermatangium, we conducted detailed observations of picking behaviour in both virgin and copulated females and compared the sperm storage conditions in the seminal receptacle between females with and without spermatangia picking after copulation in virgin females. In all observations, elongation of the buccal mass occurred within 5 min after copulation. However, sperm volume in the seminal receptacle was not related to spermatangia picking. Observations using slow-motion video revealed that females removed the spermatangia by blowing or eating after picking. These results suggest that picking behaviour is used for sperm removal but not for sperm transfer. Moreover, the frequency of buccal mass elongation was higher in copulated females than in virgin females, consistent with the sequential mate choice theory whereby virgin females secure sperm for fertilisation, while previously copulated females are more selective about their mate. Female I. paradoxus may choose its mate cryptically through postcopulatory picking behaviour