Sexually transmitted parasites and host mating behavior in the decorated cricket

Sexually transmitted diseases play a potentially important rolein the ecology and evolution of host mating behavior. Here,we use a sexually transmitted nematode-cricket (Mehdinema alii–Gryllodessigillatus) system to examine the effects of parasitism on hostmating activity and female choice. Previous work has shown thatinfected male crickets produce a significantly smaller nuptialgift (spermatophylax) than uninfected males. This is expectedto result in reduced spermatophylax feeding duration and earlyampulla removal. Here, we hypothesize that the parasite-mediatedreduction in spermatophylax size will consequently shorten femaleintercopulatory interval. We predict that females mated to infectedmales will exhibit a shorter intercopulatory interval than femalesmated to uninfected males. To test this hypothesis, we experimentallymeasured the behavioral responses of females mated to uninfectedand infected males. We found no significant difference betweenfemale handling of the spermatophylax and ampulla from infectedversus uninfected males. Although the duration of spermatophylaxconsumption is positively correlated with the duration of ampullaattachment, neither of these variables is correlated with femaleintercopulatory interval. Intercopulatory intervals for femalespreviously mated with uninfected versus infected males are notstatistically different. We conclude that parasitism in maleG. sigillatus does not influence female intercopulatory intervalor male mating success. We found no evidence for female matechoice based on male infection status. The lack of female choiceis consistent with theoretical predictions involving parasitesthat are sexually transmitted.     

营养对雄性短翅鸣螽繁殖投资的影响

螽斯交配时产生精包,精包由精护与精荚组成。交配后雌性咬食精护。吃完精护后将精荚吃掉。精护是螽斯的一种繁殖投资方式。我们研究营养对雄性螽斯投资的影响。结果表明：⑴螽斯首次交配后产生的精包、精护、精夹的鲜重以及精荚、精护干重组间差异显著,高营养组雄性投资显著高于其它组;⑵营养缺乏时,有效性比偏雌。本研究支持性差异理论以及Ｇｗｙｎｎｅ（１９９０）和Ｓｉｍｍｏｎｓ（１９９５）的工作;⑶精荚和精护重量与雄性     

Variation in nuptial gift quality in bush crickets (Orthoptera: Tettigoniidae)

Males of many insect species provide the female, during courtshipand copulation, with a nuptial gift consisting of a prey itemor synthesized material (e.g., spermatophores). The studiesperformed so far have mainly focused on effects of nuptial giftsize on male and female reproductive success. However, the qualityof the nuptial gift can differ substantially between taxa andmay potentially have a large impact on male and female reproductiveperformance. In this study the effects of. variation in dieton nuptial gift quality is investigated in several bush cricketspecies with different diets. The effect of diet on nuptialgift quality (e.g., protein in the spermatophylax) and femalereproductive output and, in turn, die effect of variation inspermatophylax quality on female reproductive output are investigated.Female reproductive output and male spermatophore size weremainly found to be influenced by differences in diet betweenspecies. Spermatophylax quality (i.e., protein concentration)was also correlated with differences in diet. There was a largevariation in protein content of the spermatophylax widiin aswell as between species. Moreover, larger spermatophylaxes hada lower protein concentration, indicating a possible trade-offbetween spermatophylax size and quality. Consequendy, productionof larger spermatophylaxes, required for protection of the male'ssperm carrying ampulla during insemination, can lead to a reducedprotein concentration, because the total amount of protein maybe limited. This pattern is also consistent with die idea diatthe spermatophylax functions primarily to ensure sperm transfer.Finally, there was no correlation between the amount of proteinin the spermatophylax and female reproductive output eitheracross diets or within each diet category, further supportingthe finding that female reproductive output is mainly affectedby differences in diet.     

Condition-dependent spermatophore size is correlated with male's age in a bushcricket (Orthoptera: Phaneropteridae)

During mating, male bushcrickets transfer a spermatophore that consists of a sperm-containing ampulla and a product of the accessory glands, the spermatophylax, which females directly ingest. In the present study, we demonstrate male spermatophore allocation in the bushcricket Poecilimon zimmeri . Males of this species show condition-dependent spermatophore investment. This investment depended upon the age at first mating of males, with older individuals transferring larger spermatophores than younger ones of the same body mass. Independently of age, heavier males transfer larger spermatophores, but the size of males (as measured by femur length) was not a good predictor. Heavier males allocate a lower proportion of their mass to spermatophores and reach their maximal investment point earlier than less heavy males. Spermatophylax production levelled off to a species specific maximum earlier than that of sperm investment (measured as ampulla mass), suggesting that males face high levels of sperm competition.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 354–360.     

Nuptial feeding by male bushcrickets: an indicator of male quality?

Male bushcrickets transfer a spermatophore at mating that consistsof a sperm-containing ampulla and a product of the accessoryglands, the spermatophylax, that is consumed by the female duringinsemination. Male Requena verticalis produce functionally differentspermatophores depending on the availability of sexually receptivefemales. They will maintain high mating frequency by providinga gift sufficient to ensure sperm transfer, or will invest parentallyin females when their mating frequency is low. We examined therelationship between male quality and nuptial feeding underconditions where males invest in ejaculate protection or inparental investment. When investing in ejaculate protection,males reduced the quality of the spermatophylax meal by reducingboth the concentration of protein and the absolute amount ofprotein it contained. There was no relationship between malephenotype and gift size or quality. Moreover, we could findno evidence for the recently advanced hypothesis that femalescan exercise mate choice by interfering with insemination. However,when males were investing parentally, we found a positive associationbetween spermatophylax size and male size, but no relationshipbetween protein content and male size. Males with high levels offluctuating asymmetry invested more heavily in the nutritionalcontent of their spermatophylaxes than did symmetrical males.Thus, male quality does influence nuptial feeding, but in amanner predicted by a model of indirect fitness benefits frommate choice.     

Pairing success of male Gammarus lacustris infected by two acanthocephalans: a comparative study

The costs of parasitism to host reproduction can be best assessedusing field studies to determine overall mating success andexperimental studies to examine how parasites may affect matingbehavior. We compared the influence of two parasites, Polymorphusparadoxus and P. marilis (Acanthocephala), on the pairing successof their intermediate host (Gammarus lacustris, Crustacea) inboth the field and laboratory. Parasitism significantly loweredthe pairing success of male gammarids. In the field, P. paradoxus-infectedmales paired significantly less often than P. marilis-infectedor uninfected males. Those infected by P. marilis were alsofound in precopula significandy less often than uninfected ones.In the laboratory, the pairing success of males infected byeither parasite was significantly reduced in both competitiveand noncompetitive situations. As in the field studies, thepairing success of P. paradoxus-infected males was significantlylower than that of P. marilis-infected and uninfected males.Polymorphus marilis-infected males were also outcompeted byuninfected individuals, however, their pairing success improvedwhen alone with a female (noncompetitive experiments). We relatethe differential influence of the two parasites on the pairingsuccess of male gammarids to their effects on the physiologyand behavior of G. lacustris.     

SPERMATOPHORE SIZE IN BUSHCRICKETS: COMPARATIVE EVIDENCE FOR NUPTIAL GIFTS AS A SPERM PROTECTION DEVICE

During courtship and copulation, males of many insect species provide the female with a nuptial gift of a prey item or synthesized material. These gifts may be explained as a form of paternal investment by increasing female reproductive output, or in terms of mating effort by increasing male fertilization success. These explanations, while not mutually exclusive, are controversial. While experimental studies examine the maintenance of nuptial gifts in single species, comparative studies are required to indicate more general evolutionary trends. Male bushcrickets provide females with a nuptial gift, a spermatophylax, which is transferred to females at mating along with the sperm-containing ampulla. Analysis of comparative data of 28 species of bushcrickets (Orthoptera: Tettigoniidae), reveals that male spermatophore size (spermatophylax and ampulla weight) is positively correlated with female refractory period, which, in turn, correlates with male fertilization success. Moreover, gift size (the spermatophylax) covaries with ejaculate size (the ampulla), which is consistent with the hypothesis that it serves as a sperm protection device. In contrast, there is no significant correlation between any measure of female fecundity and male spermatophylax size. This indicates that the variation in spermatophore size among bushcrickets is better explained by a mating-effort function than a paternal investment function.     

Cryptic female choice predicated on wing dimorphism in decorated crickets

Male decorated crickets, Gryttodes sigllatus, normally lackbind wings and are incapable of flight (short-winged males),but occasionally exhibit fully developed hind wings that makerudimentary flight possible (long-winged males). Long-wingedmales bear a cost of flight in the form of decreased inseminationsuccess, which arises as a consequence of two interrelated factors:(1) long-winged males exhibit a lower reproductive investmentrelative to short-winged males, as measured by the mass of amale's spermatophore and reproductive organs and (2) the postcopulatorybehavior of females favors males that maximize their reproductiveinvestment. Of particular importance to male mating successis the spermatophylax, a large gelatinous mass forming partof the spermatophore and consumed by the female after mating.Consumption of the spermatophylax keeps the female preoccupiedwhile sperm are discharged from the remaining portion of thespermatophore (sperm ampulla) into her repro ductive tract.The spermatophylax of long-winged males is significantly smallerthan that of short-winged males and consequently requires lesstime to consume. As a result, the sperm ampulla of long-wingedmales is frequently removed before its complete evacuation andsignificantly sooner than that of short-winged males. Becausethe spermatophore-removal behavior of females mediates the relativeinsemination success of short-winged and long-winged males,it can be considered a form of cryptic female choice     

Food fight: sexual conflict over free amino acids in the nuptial gifts of male decorated crickets

In decorated crickets, Gryllodes sigillatus, the spermatophore that a male transfers at mating includes a gelatinous spermatophylax that the female consumes, delaying her removal of the sperm‐filled ampulla. Male fertilization success increases with the length of time females spend feeding on the spermatophylax, while females may benefit by prematurely discarding the spermatophylaxes of undesirable males. This sexual conflict should favour males that produce increasingly appealing spermatophylaxes, and females that resist this manipulation. To determine the genetic basis of female spermatophylax feeding behaviour, we fed spermatophylaxes to females of nine inbred lines and found that female genotype had a major influence on spermatophylax feeding duration. The amino acid composition of the spermatophylax was also significantly heritable. There was a positive genetic correlation between spermatophylax feeding duration and the gustatory appeal of the spermatophylax. This correlation suggests that genes expressed in males that produce more manipulative spermatophylaxes are positively linked to genes expressed in females that make them more vulnerable to manipulation. Outbred females spent less time feeding on spermatophylaxes than inbred females, and thus showed greater resistance to male manipulation. Further, in a nonspermatophylax producing cricket (Acheta domesticus), females were significantly more prone to feeding on spermatophylaxes than outbred female Gryllodes. Collectively, these results suggest a history of sexually antagonistic coevolution over the consumption of nuptial food gifts.     

Postmating sexual selection: the effects of male body size and recovery period on paternity and egg production rate in a water strider

The role of male body size in postmating sexual selection wasexplored in a semiaquatic insect, the water strider Gerris lateralis.To separate effects of male size per se from those due to numericsperm competition, male recovery period (shown here to be proportionalto ejaculate size) was manipulated independently of body sizein a factorial experiment where virgin females were mated firstwith sterile males and then with focal males. Both relativemale fertilization success and female reproductive rate were measured.The number of sperm transferred increased with male recoveryperiod, an effect that was mediated by longer copulation duration,but there were no effects of body size on ejaculate size. Neithermale size nor recovery period had any significant direct effectson male fertilization success. However, copulation durationinfluenced relative fertilization success, suggesting that malesable to transfer more sperm also achieved higher fertilizationsuccess. Females exercised cryptic female choice by modulatingtheir reproductive rate in a manner favoring large males andmales that were successful in terms of achieving high relativefertilization success. Thus, successful males gained a twofoldadvantage in postmating sexual selection. This study has important implicationsfor previous estimates of sexual selection in this group of insectsbecause pre- and postmating sexual selection will be antagonisticdue to limitations in male sperm production: males mating frequently(high mating success) will on average transfer fewer sperm ineach mating and will hence tend to fertilize fewer eggs permating (low fertilization success).     

Anther-smut infection of Silene alba caused by Ustilago violacea: factors determining fungal reproduction

Summary The anther-smut fungus Ustilago violacea sporulates in flowers of the dioecious host plant Silene alba. Growth chamber comparisons of healthy and diseased plants, with the genetic background of host and pathogen controlled, revealed that fungal infection increases the number of flowers produced per plant and decreases the size of individual flowers. There were few consistent effects of plant genotype or fungal isolate on diseased flower traits, but differences between the plant sexes were apparent. Stimulation of flower production is proportionally greater in females than males: thus, although healthy male plants produce many more flowers than healthy females, sexual differences in diseased flower number are reduced. Sexual differences in diseased flower size also exist, with male flowers smaller than females. A field inoculation study confirmed dimorphism in diseased flower size and demonstrated that spore production per flower was greater for males than females for all flower size classes.     

COURTSHIP FEEDING AND THE FITNESS OF FEMALE KATYDIDS (ORTHOPTERA: TETTIGONIIDAE)

Katydid males (Requena verticalis) produce spermatophores with a large sperm-free spermatophylax, which is eaten by the female after mating. In this study, I asked 1) how do spermatophylax nutrients affect the fitness of the mated female and her progeny? 2) does this male-produced food substitute for other food in the diet of the female, or is it a source of specialized nutrients? and 3) does an increase in the size of the spermatophylax eaten influence female reproduction in the same way as the additional spermatophylaxes that would be obtained from multiple mating? An experimental increase either in dietary protein or the number of spermatophylaxes eaten increased the number of eggs produced. However, spermatophylax size had no effect on egg number. An increase in either the size or number of spermatophylaxes eaten resulted in an increase in egg size. There was no influence of protein in the general diet on egg size. This suggests that males provide nutrition not available from other sources. Although there were no direct effects of number of spermatophylaxes eaten by females on the overwintering survival of their progeny, offspring from females producing larger eggs had a relatively higher probability of surviving winter. The amount of spermatophylax eaten had no influence on the mean adult size of progeny but significantly increased the mean date at which sons matured to adulthood. There was no influence of dietary protein on these variables. Since maturation date is positively correlated with adult size in both sexes, it is suggested that the influence of courtship feeding on maturation date may result in an increase in adult size and thus the fitness of sons. A significant correlation between the size of the female and the mean size of her sons (but not daughters) suggests that there is also a heritable component to body size.     

Ejaculate expenditure by malebush crickets decreases with sperm competition intensity

Male bushcrickets transfer a spermatophore at mating that consists of a sperm-containing ampulla and a sperm-free mass, the spermatophylax, that is consumed by the female during insemination. The costs of spermatophore production for males and benefits of consumption for females result in reversals in courtship roles in nutrient limited populations that increase both the risk and intensity of sperm competition. Here we show that under conditions characteristic of courtship role reversal, male expenditure on the spermatophore is dependent on female size. When mating with small females, males increase the amount of spermatophylax material and sperm, as expected from the increased sperm competition risk associated with courtship role reversal. However, males reduce the amount of spermatophylax material and sperm transferred to larger females. Since larger females have a higher mating success when competing for nurturant males, the intensity of sperm competition covaries with female size. Reduced ejaculate expenditure under increased sperm competition intensity is in accord with theoretical expectation.     

Paternity of offspring in multiply-mated,female crickets: the effect of nuptial food gifts and the advantage of mating first

The spermatophore transferred by male decorated crickets (Gryllodes sigillatus) includes a large gelatinous mass, the spermatophylax, that is consumed by the female after mating. This nuptial gift preoccupies the female while sperm are discharged from the remaining portion of the spermatophore, the sperm ampulla, into her reproductive tract. There is considerable variation in the mass of the spermatophylax, and about half of all males produce spermatophylaxes that are too small to ensure complete sperm transfer. We tested two hypotheses concerning the maintenance of this variation: (i) males trade-off investment in spermatophylaxes against copulation frequency; and (ii) males synthesize the largest spermatophylaxes of which they are physiologically capable. Males synthesizing large and small food gifts were permitted multiple mating opportunities with the same females, and allozyme markers were used to establish the paternity of offspring. There was a significant advantage to those males that mated first irrespective of gift size. This advantage probably arose, in part, because the sperm of first males would have had exclusive access to females'' eggs during the first 24 hours of oviposition, and underscores the benefits of matings with virgin females. The paternity of ''small-gift'' males increased with gift mass, but there was no such increase in ''large-gift'' males. This difference probably stems from the relationship between gift mass and sperm transfer: most of the gifts of the large-gift males would have been above the threshold needed to achieve complete inseminations, whereas those of small-gift males would have been below the threshold. Within mating-order positions, there was no significant difference in the paternity of large-gift and small-gift males, a result seemingly consistent with the ''trade-off'' hypothesis. However, there was no correlation between spermatophylax mass and male mating frequency, so that the mechanism by which small-gift males offset their fertilization disadvantage remains unknown.     

Male-biased sex-ratio distortion caused by Octosporea bayeri, a vertically and horizontally-transmitted parasite of Daphnia magna

Female-biased sex-ratio distortion is often observed in hosts infected with vertically-transmitted microsporidian parasites. This bias is assumed to benefit the spread of the parasite, because male offspring usually do not transmit the parasite further. The present study reports on sex-ratio distortion in a host-parasite system with both horizontal and vertical parasite transmission: the microsporidium Octosporea bayeri and its host, the planktonic cladoceran Daphnia magna. In laboratory and field experiments, we found an overall higher proportion of male offspring in infected than in uninfected hosts. In young males, there was no parasite effect on sperm production, but, later in life, infected males produced significantly less sperm than uninfected controls. This shows that infected males are fertile. As males are unlikely to transmit the parasite vertically, an increase in male production could be advantageous to the host during phases of sexual reproduction, because infected mothers may obtain uninfected grandchildren through their sons. Life-table experiments showed that, overall, sons harboured more parasite spores than their sisters, although they reached a smaller body size and died earlier. Male production may thus be beneficial for the parasite when horizontal transmission has a large pay-off as males may contribute more effectively to parasite spread than females.     

Sex allocation and effects of superparasitism on secondary sex ratios in the gregarious parasitoid, Trichogramma chilonis (Hymenoptera: Trichogrammatidae)

The role of sex-controlling behaviour at oviposition in generating primary sex ratios, and the effect of larval competition on secondary sex ratios, were studied in the gregarious endoparasitoid, Trichogramma chilonis. The production of a fertilized (female) egg is indicated by the incorporation of a pause in abdominal movements during oviposition, while the absence of it indicates the production of an unfertilized (male) egg. During each ovipositional bout, the first male egg is deposited at the second oviposition, and thereafter at intervals of about eight eggs. This simple pattern enables the wasps to adjust their progeny sex ratios under local male competition to a wide range of host size. Inexperienced wasps do not distinguish between parasitized and healthy hosts. Immature mortality is not significantly different between the sexes when a host is attacked by a single wasp, while females suffer higher immature mortality than males when superparasitism occurs.     

Identifying a causal agent of sexual selection on weaponry in an insect

In many animal species, males do not seek females directly butinstead locate and defend sites that contain spatially or temporallylimited resources essential to female survival and reproduction.Resident males that successfully repel conspecific rivals canmate with females attracted to these resources. In theory, increasingresource value increases harem size and thus increases the opportunity(I_mates) for and strength of sexual selection on traits crucialto male resource-holding potential and mating success. I experimentallytested this hypothesis in the field using the Wellington treeweta, Hemideina crassidens (Orthoptera: Tettigonioidea: Anostostomatidae),a sexually dimorphic insect in which males use their enlargedmandibles as weapons in male–male contests over accessto females sheltering in tree cavities (galleries). By manipulatinggallery size, I showed that, compared with smaller galleries,larger galleries housed larger harems. Variation in gallerysize was an important determinant of I_mates, but contrary toexpectation, greater opportunity existed in small galleriescompared with large galleries. As predicted, male weapon sizewas under stronger directional selection in large galleriesbecause the fitness benefits were greater under these conditionscompared with small galleries. My results help explain the positiveassociation between average weapon size and average gallerysize observed within and among tree weta populations in NewZealand.     

Mortality effects of the parasitoid flesh fly Blaesoxipha japonensis (Diptera: Sarcophagidae) in relation to body size of the adult grasshopper Parapodisma tanbaensis (Orthoptera: Catantopidae)

Body size is a significant factor for organisms because it has a profound effect on almost all of their physiological, life-history, ecological, and evolutionary traits. Sexual size dimorphism (SSD) is one of the sources creating intraspecific variation in body size. In general, female adults are larger than male adults in insects. Whereas much research has examined the significance of male competition and fecundity as causes of SSD in insects, relatively less is known about the effects of parasitoids on host insect size. To investigate the effects of the parasitoid fly Blaesoxipha japonensis Hori on the body size of the adult grasshopper Parapodisma tanbaensis Tominaga et Kanô, we examined the relationship between adult host size and host mortality rate caused by B. japonensis when adult grasshoppers captured in the field were reared in the laboratory. The results showed that females were larger than males and that the host mortality rate caused by B. japonensis was inversely related to host size in females, whereas there was no significant relationship between male size and host mortality rate. These results suggest that the effect of this parasitoid on host body size is one factor favoring the female-biased SSD in this adult grasshopper.     

Colony pollen reserves affect body size,sperm production and sexual development in males of the stingless bee <Emphasis Type="Italic">Melipona beecheii</Emphasis>

The production of male sexual offspring by social insect colonies is often strongly seasonal or resource-dependent. In stingless bees, males are produced in smaller numbers under conditions of low colony food reserves; whether such males are negatively affected in traits related to reproductive success is not known. We compared body size, sperm production and sexual maturity in Melipona beecheii males reared with experimentally supplemented or reduced pollen reserves, but with otherwise equal numbers of workers and equal quantities of honey reserves. We also studied the same traits in males collected from non-manipulated colonies with pollen reserves intermediate between the supplemented or reduced groups but with more workers and honey reserves. Males reared under experimentally reduced pollen reserves had significantly smaller bodies and lower sperm counts compared to those reared in colonies with experimentally supplemented pollen reserves. There was also a significantly positive relationship between the number of sperm and body size in males across all colony treatments. The maximum number of sperm in seminal vesicles was recorded 2 days later in males from colonies with reduced pollen compared to males from colonies with supplementary pollen. Males from non-manipulated colonies were intermediate in size, sperm count and speed of maturation. Our study documents for the first time the existence of large size variation in males of stingless bees that is related with the amount of pollen reserves in their natal colony. We conclude that a colony’s pollen reserves have a major impact on male body size, sperm production and speed of sexual maturity in this stingless bee, which may be the case in other social insects. Stingless bees are a good model system to study the balance between colony-level selection and individual-level selection on male sexually selected traits such as body size.