营养质量对雄性赤拟谷盗生长发育、抗饥饿和资源再获取能力的影响

昆虫的生长发育、抗饥饿和资源再获取能力对其生存、生殖和性选择非常重要。本文研究了营养质量对赤拟谷盗Tribolium castaneum(Herbst)生长发育、抗饥饿和资源再获取能力的影响。结果表明,三个营养处理下雄性赤拟谷盗的生长发育性状（幼虫重、发育历期、蛹重和成虫重）均存在显著性差异,在高营养质量下生长的雄性赤拟谷盗发育更快,体重更大,且各生长发育性状间存在显著性相关。营养质量对雄性赤拟谷盗成虫的抗饥饿能力没有显著影响,但对其资源再获取能力有显著影响,表现为在高营养质量下生长的雄性赤拟谷盗成虫比在低营养质量下生长的具有更高的资源再获取能力。雄性赤拟谷盗抗饥饿能力和资源再获取能力之间存在显著性相关,但是雄性赤拟谷盗抗饥饿能力和资源再获取能力除了与发育历期存在显著性相关外,与其它生长发育性状不存在显著性相关。本文还对这些实验结果的性选择意义进行了讨论。     

Genetic and nutritional effects on male traits and reproductive performance in Tribolium flour beetles

In Tribolium flour beetles and other organisms, individuals migrate between heterogeneous environments where they often encounter markedly different nutritional conditions. Under these circumstances, theory suggests that genotype-by-environment interactions (GEI) may be important in facilitating adaptation to new environments and maintaining genetic variation for male traits subject to directional selection. Here, we used a nested half-sib breeding design with Tribolium castaneum to partition the separate and joint effects of male genotype and nutritional environment on phenotypic variation in a comprehensive suite of life-history traits, reproductive performance measures across three sequential sexual selection episodes, and fitness. When male genotypes were tested across three nutritional environments, considerable phenotypic plasticity was found for male mating and insemination success, longevity and traits related to larval development. Our results also revealed significant additive genetic variation for male mating rate, sperm offence ability (P(2)), longevity and total fitness and for several traits reflecting both larval and adult resource use. In addition, we found evidence supporting GEI for sperm defence ability (P(1)), adult longevity and larval development; thus, no single male genotype outperforms others in every nutritional environment. These results provide insight into the potential roles of phenotypic plasticity and GEI in facilitating Tribolium adaptation to new environments in ecological and evolutionary time.     

The influence of the nutritional content of the adult male diet on testis mass, body condition and courtship vigour in a Hawaiian Drosophila

1. The effects of the nutritional quality of the adult diet (primarily protein content) on testis mass, body condition and courtship vigour were studied in a Hawaiian Drosophila, D. grimshawi , a lek-forming species under strong sexual selection. The primary goals of this study were to determine whether there is a trade-off between investment in reproductive and somatic tissues, and to examine whether this trade-off is influenced by quality of the adult diet.
2. Quality of the adult diet had a major influence on male body condition, courtship vigour and testis mass, but males varied in their investment patterns even within diet treatment.
3. Body condition, a measure of phenotypic quality, was significantly related to how much males invested in testis and body tissues, but the nature of the relationships differed between males fed high- or low-quality diets.
4. Paragonia volume was significantly smaller for adult males fed low-quality diets than for males fed high-quality diets. Nutrient-deficient diets apparently forced males to trade off investment in testes and body condition, and in body condition and courtship vigour, but nutrient-rich diets did not result in severe trade-offs.
5. Collectively, the results suggest that fluctuation in adult diet quality of male D. grimshawi might influence male reproductive quality in ways that might limit female reproductive output and/or be a factor for female choice in this species.     

Male field crickets that provide reproductive benefits to females incur higher costs

Abstract.  1. Females often select mates based on signals correlated with the quality of the direct benefits that males will provide to them. A male's quality as a mate and the structure of his mating signals may covary because both traits are energetically expensive for males to produce and because both traits are affected by short-term changes in nutritional condition.
2. In the variable field cricket, Gryllus lineaticeps , previous work has shown that females receive reproductive benefits from males that produce higher chirp rates and lifespan benefits from males that produce longer chirp durations, even when they only receive the sperm and seminal fluid contained in male spermatophores. Higher chirp rates are energetically expensive for males to produce, and chirp rate is strongly affected by diet quality, whereas longer chirp durations do not appear to be expensive for males to produce, and chirp duration does not appear to be affected by male diet quality. In this study two hypotheses were tested about the energetic costs of spermatophore production: (1) that spermatophores are expensive for males to produce and (2) that males providing greater direct benefits to females incur higher costs of spermatophore production.
3. Males that were provided with a lower quality diet took longer to produce a new spermatophore. This result suggests that spermatophores are costly for males to produce.
4. Males that produced higher chirp rates took longer to produce a new spermatophore. This result suggests that male chirp rate and female reproductive benefits may covary because both traits are energetically expensive for males to produce and thus are affected by male nutritional condition. There was no association, however, between male chirp duration and spermatophore production time.     

Pheromone production by male Tribolium castaneum (Coleoptera: Tenebrionidae) is influenced by diet quality

Tribolium castaneum (Herbst) (Coleoptera: Tenebrionidae), the red flour beetle, is a common cosmopolitan pest exploiting a variety of stored products. We experimentally manipulated diet nutritional quality by using non-nutritive filler to examine how this influenced pheromone production and olfactory attractiveness of T. castaneum adult males. Volatiles released by individual males reared on high versus low nutrition diets were collected using solid phase microextraction, and gas chromatography coupled to mass spectrometry was used to identify and quantify the Tribolium aggregation pheromone 4, 8-dimethyldecanal (DMD). Males kept on high nutrition diet showed a three-fold increase in daily DMD production, which suggests the possibility that this pheromone could act as a condition-dependent mating signal. In pitfall trap assays, there was no significant difference in the mean response of virgin females to discs kept with low versus high nutrition males, although discs carrying male cues were significantly more attractive than blank discs. These results suggest that DMD production rates by T. castaneum males will depend on the nutritional quality of various stored products, but such differences may not alter males' ability to attract females.     

Examining the potential for resource-dependent female reproductive fluid-sperm interactive effects in a livebearing fish

Sexually selected traits can be costly to produce and maintain. The amount of resources available to an individual is therefore expected to influence investment in costly sexual traits. While resource-dependent expression of sexually selected traits has traditionally been examined in males, resource limitation can also influence how sexual selection operates in females. Female reproductive fluids are thought to be costly to produce and may play an important role in shaping the outcome of postcopulatory sexual selection by influencing sperm performance. However, we know surprisingly little about whether and how female reproductive fluids are influenced by resource limitation. Here, we examine if resource restriction influences female reproductive fluid-sperm interactive effects in the pygmy halfbeak (Dermogenys collettei), a small internally fertilizing freshwater fish where females store sperm. After experimentally altering female diets (high vs. restricted diets), we compared how female reproductive fluids influence two key metrics of sperm quality: sperm viability and velocity. While female reproductive fluids enhanced sperm viability and velocity, we found no evidence that female diet influenced the interactive effect between female reproductive fluids and sperm viability or velocity. Our findings build on the growing evidence that female reproductive fluids influence sperm performance and call for further attention to be devoted to understanding how resource quantity and quality influence how female reproductive fluids affect sperm performance.     

Intraspecific variation in ejaculate traits of the northern watersnake (Nerodia sipedon)

Sperm competition occurs when the sperm of more than one male compete to fertilize the eggs of a female. In reptiles, sperm competition is particularly prevalent and is an important agent of sexual selection in males. Spermatogenesis in reptiles can be energetically expensive, suggesting that there may be costs to producing high-quality ejaculates. The northern watersnake Nerodia sipedon has a mating system characterized by aggregations in which a single female mates with multiple males, resulting in high levels of multiple paternities. Under these circumstances sperm competition is likely important, and selection should favour sperm and ejaculate traits that enhance a male's reproductive success. In this study, we examined intraspecific variation in ejaculate quality (sperm length, motility, sperm density, spermactocrit) in male watersnakes and determined whether ejaculate traits varied with body size and condition, using both size-corrected mass and haematocrit as indices of condition. We found large variation among males in all these traits, except for sperm length. Although there was significant variation in sperm length among males, the majority of variation in sperm length occurred within rather than among individuals. Males with high haematocrit had sperm that were less variable with respect to length, and large males produced ejaculates that were less concentrated with respect to sperm than small males. The lack of condition dependence of most ejaculate traits is consistent with previous studies that indicate that male reproductive effort in this species is generally not energy limited, perhaps because of opportunistic foraging during the mating season.     

Female life span and fertility are increased by the ejaculates of preferred males

In animals with internal fertilization, sperm competition among males can favor the evolution of male ejaculate traits that are detrimental to females. Female mating preferences, in contrast, often favor traits in males that are beneficial to females, yet little is known about the effect of these preferences on the evolution of male ejaculates. A necessary condition for female preferences to affect the evolution of male ejaculate characteristics is that females select mates based on a trait correlated with ejaculate quality. Previous work has shown that females of the variable field cricket, Gryllus lineaticeps, prefer males that produce calling songs containing faster and longer chirps. In this study, we tested the hypothesis that females receive more beneficial ejaculates from preferred males. Females were placed on either a high- or a reduced-nutrition diet then mated twice to a male of known song phenotype. Females received only sperm and seminal fluid from males during these matings. There was no effect of male song phenotype on any fitness component for females on the high-nutrition diet. Reduced-nutrition females mated to males that produced preferred song types, however, lived longer, produced more eggs, produced more fertile eggs, and had a higher proportion of their eggs fertilized than those mated to other males. The life-span benefit was positively associated with male chirp duration, and the reproductive benefits were positively associated with male chirp rate. We explored two possible mechanisms for the life span and reproductive benefits. First, a path analysis suggested that part of the effect of male chirp duration on female life span may have been indirect; females mated to males that produced longer chirps showed delayed oviposition, and females that delayed oviposition lived longer. Males that produce longer chirps may thus transfer fewer or less potent oviposition stimulants to females in their seminal fluid. Second, there was a positive correlation between male chirp rate and the number of sperm transferred to females. The fertility benefit may thus have resulted from females receiving more sperm from males that produce faster chirps. Finally, there was a negative phenotypic correlation between male chirp rate and chirp duration, suggesting that females may have to trade off the life span and reproduction benefits when selecting a mate.     

Resource quality affects weapon and testis size and the ability of these traits to respond to selection in the leaf‐footed cactus bug,Narnia femorata

The size of weapons and testes can be central to male reproductive success. Yet, the expression of these traits is often extremely variable. Studies are needed that take a more complete organism perspective, investigating the sources of variation in both traits simultaneously and using developmental conditions that mimic those in nature. In this study, we investigated the components of variation in weapon and testis sizes using the leaf‐footed cactus bug, Narnia femorata (Hemiptera: Coreidae) on three natural developmental diets. We show that the developmental diet has profound effects on both weapon and testis expression and scaling. Intriguingly, males in the medium‐quality diet express large weapons but have relatively tiny testes, suggesting complex allocation decisions. We also find that heritability, evolvability, and additive genetic variation are highest in the high‐quality diet for testis and body mass. This result suggests that these traits may have an enhanced ability to respond to selection during a small window of time each year when this diet is available. Taken together, these results illustrate that normal, seasonal fluctuations in the nutritional environment may play a large role in the expression of sexually selected traits and the ability of these traits to respond to selection.     

Adult male nutrition and reproductive success in Drosophila melanogaster

Explanations for the maintenance of variation in reproductive traits influenced by seminal fluid accessory gland proteins (Acps) in male Drosophila melanogaster include nontransitivity in the outcome of sperm competition and/or condition dependence of the traits involved. We investigated the effects of adult male nutrition (five diets) on the expression of Acp- and sperm- mediated traits. We found novel, nonlinear effects, with females showing lower levels of refractoriness to remating after mating with males held on the lowest and highest yeast diets. There were no significant effects of adult male nutrition on male paternity share, but there was a striking, nonlinear effect on second male progeny production, with males kept on intermediate yeast diets fathering the highest number of offspring. Such "bell shaped" responses of life-history traits to nutrition have only previously been reported for longevity. Consistent with previous reports, males maintained on low protein diets had lower premating success and gained fewer rematings with nonvirgins. We show novel and body size independent effects of adult male nutrition on traits influenced by Acps and sperm, which do not fit current condition-dependent handicap models and can affect the strength of sexual selection acting upon such fitness-related traits.     

An integrative view of sexual selection in Tribolium flour beetles

Sexual selection is a major force driving the evolution of diverse reproductive traits. This evolutionary process is based on individual reproductive advantages that arise either through intrasexual competition or through intersexual choice and conflict. While classical studies of sexual selection focused mainly on differences in male mating success, more recent work has focused on the differences in paternity share that may arise through sperm competition or cryptic female choice whenever females mate with multiple males. Thus, an integrative view of sexual selection needs to encompass processes that occur not only before copulation (pre-mating), but also during copulation (peri-mating), as well as after copulation (post-mating), all of which can generate differences in reproductive success. By encompassing mechanisms of sexual selection across all of these sequential reproductive stages this review takes an integrative approach to sexual selection in Tribolium flour beetles (Coleoptera: Tenebrionidae), a particularly well-studied and economically important model organism. Tribolium flour beetles colonize patchily distributed grain stores, and juvenile and adult stages share the same food resources. Adults are highly promiscuous and female reproduction is distributed across an adult lifespan lasting approximately 1 year. While Tribolium males produce an aggregation pheromone that attracts both sexes, there appears to be little pre-mating discrimination among potential mates by either sex. However, recent work has revealed several peri-mating and post-mating mechanisms that determine how offspring paternity is apportioned among a female's mates. During mating, Tribolium females reject spermatophore transfer and limit sperm numbers transferred by males with low phenotypic quality. Although there is some conflicting evidence, male copulatory leg-rubbing appears to be associated with overcoming female resistance to insemination and does not influence a male's subsequent paternity share. Evidence suggests that Tribolium beetles have several possible post-mating mechanisms that they may use to bias paternity. Male sperm precedence has been extensively studied in Tribolium spp. and the related Tenebrio molitor, and several factors influencing male paternity share among a female's progeny have been identified. These include oviposition time, inter-mating interval, male strain/genotype, the mating regimen of a male's mother, male starvation, and tapeworm infection. Females exert muscular control over sperm storage, although there is no evidence to date that females use this to differentiate among mates. Females could also influence offspring paternity by re-mating with additional males, and T. castaneum females more readily accept spermatophores when they are re-mating with more attractive males. Additional work is needed to examine the possible roles played by both male and female accessory gland products in determining male paternity share. Sexual selection during pre-mating episodes may be reinforced or counteracted by peri- and post-copulatory selection, and antagonistic coevolution between the sexes may be played out across reproductive stages. In Tribolium, males' olfactory attractiveness is positively correlated with both insemination success and paternity share, suggesting consistent selection across different reproductive stages. Similar studies across sequential reproductive stages are needed in other taxa to provide a more integrative view of sexual selection.     

Trade-off between pre and post-copulatory traits depends on locomotor activity in male Tribolium castaneum beetles

Locomotor performance is an indicator of dynamic exercise; thus, it is a central trait in many animal behaviours. Although higher locomotor endurance may increase male reproductive success (e.g., in mate searching and male–male contests), investment in other male reproductive traits (e.g., male attractiveness and sperm competition) may be decreased through energy consumption due to higher activity levels. Here, I investigated male attractiveness, mating success, and paternity success using males of the red flour beetle Tribolium castaneum selected for higher (H) and lower (L) locomotor endurance. Although there was no difference in male attractiveness between the selection regimes, H males had significantly higher mating success than L males. Conversely, L males had significantly higher paternity success than H males. Therefore, there was a trade-off between mating success and paternity success among the selection regimes, suggesting that locomotor endurance affects male reproduction in T. castaneum, and individual variation of locomotor endurance may be maintained within a population.     

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 cloned-cattle meat on reproductive physiology in rats

The influence of the cloned-cattle meat diets upon reproduction in mammals was rarely studied. This study was performed to analyze the effects of the diets containing cloned-cattle (Korean native beef, Hanwoo) meat on the reproductive physiology in rats. The male and female rats were fed with the diets containing 5% or 10% of normal- (N-5 or N-10) or cloned- (C-5 or C-10) cattle meat during test periods. The rats fed with commercial pellets were used as control. Lower food consumption in normal- and cloned-cattle meat diet groups is detected in both male and female rats compared with that of control (P < 0.05, 0.01 and 0.001). No signs of cloned-cattle meat diets on male reproductive parameters are found in all groups, except for lower sperm deformity in C-5 group (P < 0.05) and higher testosterone concentration in C-10 group (P < 0.05), respectively. There are no significant test substance-related differences of Caesarean section and delivery in dams and external examination and physiological development test in neonate compared with control and normal meat groups. Based on these results, it can be postulated that there are no obvious negative effects on the reproductive physiology in rats fed with cloned-cattle meat diets compared to their comparators.     

Effects of dietary astaxanthin supplementation on reproductive characteristics of rainbow trout (Oncorhynchus mykiss)

The effect of dietary astaxanthin supplementation on reproductive characteristics was investigated in five groups of female rainbow trout broodstock fed diets containing either 0.07, 12.46, 33.33, 65.06 or 92.91 mg astaxanthin kg^?1, respectively, and two groups of male rainbow trout broodstock fed diets supplemented with 0.07 and 33.33 mg astaxanthin kg^?1, respectively, for 6 months in an artificial photoperiod system until sexual maturation. The eggs from each group of female broodstock were divided into two equal batches. One batch was fertilized with homogenized sperm of four males fed diets with 0.07 mg astaxanthin kg^?1 and the other portion with sperm of four males fed diets with 33.3 mg astaxanthin kg^?1. The females produced eggs with astaxanthin concentrations ranging from 2.03 to 29.79 mg kg^?1. Dietary astaxanthin supplementation had positive effects on investigated reproductive traits. Significant differences in rate of fertilization, percentage of eyed and hatched eggs, and mortality of eyed eggs were found between treatments (P < 0.05), but no significant difference was found on percentage of mortality before hatching (P > 0.05). A significant difference (P < 0.05) in fertilization rate was found for male groups fed 0.07 and 33.3 mg astaxanthin kg^?1. The astaxanthin content in the eggs and fertilization rate, eyed‐egg percentage and percentage hatch were significantly correlated (P < 0.05). It is concluded that dietary supplements of astaxanthin are required for optimum reproduction in rainbow trout.     

Condition‐dependent ejaculate production affects male mating behavior in the common bedbug Cimex lectularius

Food availability in the environment is often low and variable, constraining organisms in their resource allocation to different life‐history traits. For example, variation in food availability is likely to induce condition‐dependent investment in reproduction. Further, diet has been shown to affect ejaculate size, composition and quality. How these effects translate into male reproductive success or change male mating behavior is still largely unknown. Here, we concentrated on the effect of meal size on ejaculate production, male reproductive success and mating behavior in the common bedbug Cimex lectularius. We analyzed the production of sperm and seminal fluid within three different feeding regimes in six different populations. Males receiving large meals produced significantly more sperm and seminal fluid than males receiving small meals or no meals at all. While such condition‐dependent ejaculate production did not affect the number of offspring produced after a single mating, food‐restricted males could perform significantly fewer matings than fully fed males. Therefore, in a multiple mating context food‐restricted males paid a fitness cost and might have to adjust their mating strategy according to the ejaculate available to them. Our results indicate that meal size has no direct effect on ejaculate quality, but food availability forces a condition‐dependent mating rate on males. Environmental variation translating into variation in male reproductive traits reveals that natural selection can interact with sexual selection and shape reproductive traits. As males can modulate their ejaculate size depending on the mating situation, future studies are needed to elucidate whether environmental variation affecting the amount of ejaculate available might induce different mating strategies.     

Relationships among male sexually selected traits in the bean bug,Riptortus pedestris (Heteroptera: Alydidae)

Current concepts of sexual selection suggest that male reproductive success is determined by multiple sexual traits. As expression and production of multiple sexual traits are frequently associated with each other, positive or negative correlations among multiple sexual traits ensue. These relationships among traits associated with male reproductive success may be crucial in the evolution of male reproductive strategies. Here, we investigate phenotypic relationships among sexually selected traits in the armed bean bug Riptortus pedestris. In this insect, males with a larger body and weapon are more likely to win male–male competitions, and males with a larger weapon or higher courtship rate are more attractive to females. There was a significant positive correlation between body size and weapon size, whereas the courtship rate had significant negative correlations with body size and weapon size. Our results suggested that there was a phenotypic trade‐off between courtship rate and male morphology. In this insect, smaller males may make more effort in courtship behavior as an alternative mating tactic.     

Correlated responses to selection on female egg size in male reproductive traits in a butterfly

Genetic correlations between male and female traits can act as evolutionary constraints and, if involving reproductive traits, potentially influence sexual selection. Artificial selection on egg size in the tropical butterfly Bicyclus anynana has yielded highly divergent lines. Here we report evidence for correlated evolution in male traits. Males from the large-egg selected lines produced significantly heavier spermatophores independent of body size and tended to have more fertile sperm stored in their reproductive tracts than those from the small-egg selected lines. This may be due to an underlying genetic correlation in reproductive effort between the sexes. However, non-fertile sperm number and testis size remained unaffected by selection on egg size. Phenotypic correlations within an unselected population revealed that spermatophore mass and fertile sperm number, but not testis size and non-fertile sperm number, were positively related to male body size, and that larger spermatophores contained more fertile, but not non-fertile sperm. In addition, males provided larger females with bigger spermatophores and more fertile sperm, indicating males may be exercising mate choice during copulation.     

The effects of male social environment on sperm phenotype and genome integrity

Sperm function and quality are primary determinants of male reproductive performance and hence fitness. The presence of rival males has been shown to affect ejaculate and sperm traits in a wide range of taxa. However, male physiological conditions may not only affect sperm phenotypic traits but also their genetic and epigenetic signatures, affecting the fitness of the resulting offspring. We investigated the effects of male‐male competition on sperm quality using TUNEL assays and geometric morphometrics in the zebrafish, Danio rerio. We found that the sperm produced by males exposed to high male–male competition had smaller heads but larger midpiece and flagellum than sperm produced by males under low competition. Head and flagella also appeared less sensitive to the osmotic stress induced by activation with water. In addition, more sperm showed signals of DNA damage in ejaculates of males under high competition. These findings suggest that the presence of a rival male may have positive effects on sperm phenotypic traits but negative effects on sperm DNA integrity. Overall, males facing the presence of rival males may produce faster swimming and more competitive sperm but this may come at a cost for the next generation.     

Variation in plasma steroids and reproductive traits in dimorphic males of corkwing wrasse (Symphodus melops L.)

Differences in plasma steroid levels may reflect behavioral, gonadal, or morphological specializations that characterize different male reproductive phenotypes in teleost fishes. In the present study, we investigated whether differences in plasma steroid levels exist between two distinct male morphs shown in the corkwing wrasse Symphodus melops. In addition, we examined differences in male reproductive traits, including gonad mass, sperm motility, and sperm concentration, between the two male types. Possible associations between plasma steroid levels and reproductive traits were also investigated. The results indicated that males with typical male secondary sexual characters (territorial males) had higher plasma levels of 11-ketotestosterone (11kT) compared to smaller males with female secondary sexual characters (female mimics). The female mimics in turn had higher plasma levels of both testosterone (T) and 17beta-estradiol. In addition, female mimics had relatively larger gonads and longer-lived sperm than territorial males. Relative gonad mass covaried significantly with the plasma levels of 11kT and T among the morphs, indicating that the relative gonad size correlates positively with the plasma level of these two steroids. However, there was no significant covariance between sperm traits and plasma steroids for the two male morphs. Hence, our results do not indicate any causal link between sperm quality and hormones.