Effect of diet and mating status on ovarian development and oviposition in the polyphagous predator Macrolophus caliginosus (Heteroptera: Miridae)

The mirid bug Macrolophus caliginosus is commercially reared on eggs of Ephestia kuehniella, constituting an effective but expensive factitious food. Artificial diets can decrease the rearing costs of this natural enemy, but developing and evaluating an artificial diet is a very time-consuming activity. In the current study, development and reproduction of M. caliginosus on two artificial diets based on egg yolk were investigated. The artificial diets resulted in longer development and lower adult weights, but survival was comparable with that of control insects fed E. kuehniella eggs. Reproductive potential of the predator reared on factitious and artificial foods was assessed using a dissection method. The influence of nymphal food on fecundity was less important than that of adult food. Adults fed E. kuehniella eggs had a preoviposition period of about 4 days, whereas adults offered only plant material started laying eggs about 7 days after emergence. Ovarian scores at day 7 were higher for females fed E. kuehniella eggs than for those given access only to a tobacco leaf. Ovarian scores were not significantly affected by mating status. In a final test, a parallel comparison of two methods for assessing reproductive response to diet was made. Here, adult couples were offered one of four diets: E. kuehniella eggs, one of two artificial diets or no food. Half of the females were dissected and the other half was held for determining lifetime oviposition. Females fed E. kuehniella eggs had superior ovarian scores and laid more eggs than those fed either artificial diet or those given no extra food. A good correlation (r = 0.97) was obtained between ovarian scores and oviposition data, indicating that dissecting females after 1 week provides a reliable estimate of fecundity as affected by diet quality. Rapid reproductive assessments as used in the current study will help to increase the rate of development of artificial diets and may contribute to more cost effective production methods for augmentative biological control agents.     

What determines the number of ovarioles in a fly ovary?

The relation between the size of a fly and the number of ovarioles in its ovary was investigated in Phormia regina and Sarcophaga bullata. Small flies of varying size were produced by taking larvae prematurely off the food. The smallest flies thus obtained were derived from larvae only $\text{1}\text{8}$ of the weight of a normal larva. The number of ovarioles in an ovary is directly proportional to the size of the fly and, in the extreme case, is about $\text{1}\text{5}$ the normal number in Sarcophaga and about $\text{1}\text{3}$ in Phormia. Larvae prematurely taken off the food, but fed again after starving for several days, grow to normal or almost normal size and develop ovaries with about the normal number of ovarioles. Small or re-fed Sarcophaga do not show any changes in the anatomy of individual ovarioles but in Phormia disorders in ovariole development and a consequent reduction of fertility are frequent. The number of ovarioles remains identical from the early pupal stage all through the development of the pharate fly and then through ovarian development in the adult fly: it is determined by the size of the larva when it was taken off the food. This shows that it is not lack of space in a small adult fly abdomen which determines the number nor the occurrence of degenerative processes during ovarian development.     

Effects of the juvenile hormone analogue methoprene and dietary protein on male melon fly Bactrocera cucurbitae (Diptera: Tephritidae) mating success

The effect of access to dietary protein (P) and the topical application of a juvenile hormone analogue (methoprene (M)) on mating behaviour of male melon fly Bactrocera cucurbitae was assessed in the laboratory and in field cages. Age, dietary protein and methoprene application increased the mating success and influenced the mating behaviour. Treatment with methoprene (M+) to protein-deprived (P−) males had only a modest effect on the acceleration of sexual maturity, but application of methoprene (M+) to protein-fed (P+) males greatly accelerated sexual maturity. Protein diet (P+) increased mating success of males in comparison to protein-deprived (P−) males. Protein and methoprene have a synergistic effect on mating behaviour, since M + P+ treated males exhibit reduced mating latency and achieved higher mating in younger ages than methoprene and/or protein-deprived males. Copulation duration was correlated with nutritional status and M + P+ males copulated longer at the age of advanced sexual maturity than M − P+ males. Our results suggest that in this species with a lek mating system, females discriminate between the males based on their sexual signals, which were influenced by protein in the adult diet, methoprene application and age. The results are discussed in the light of mating competitiveness of precocious treated young males and their relevance to Sterile Insect Technique application against this pest species.     

Sex differences in movement between natural feeding and mating sites and tradeoffs between food consumption,mating success and predator evasion in Mediterranean fruit flies (Diptera: Tephritidae)

Summary Systematic quantitative observations of the location and diel pattern of adult Mediterranean fruit fly, Ceratitis capitata (Wiedemann), activities were carried out in an orange grove and surroundings on the island of Chios in Greece. Natural fly foods were assessed for their contribution to fly longevity, fecundity and fertility. There were diel shifts in male and female location. Females required a substantial and varied diet to realize peak fecundity. This diet was acquired away from the primary host, orange. Foraging for food throughout most of the day on fig and non-host foliage (including feeding on bird droppings) as well as on fig fruit and grapes, females dispersed and fed more than males. A diet of grapes alone did not support any fecundity, contributing only to longevity. A diet of figs alone, on the other hand, sustained both longevity and egg production. Bird feces alone supported neither egg production nor longevity. However, when added to a diet of figs, bird feces significantly increased fly fecundity. Throughout most of the day, males aggregated in leks within the inner canopy of the primary host, orange. The arrival here during the warmest hours of the day of receptive females, followed by pair formation, reinforced the lek mating system on host foliage. In the afternoon, females shifted to orange fruit where they suffered from high predation mortality while ovipositing. Soon after, males also shifted to orange fruit, where they attempted matings with non-receptive ovipositing females. Male feeding on fig fruit occurred late in the day, a time when they were least likely to find a mate. Male survival did not differ between the natural diets. Tradeoffs between food consumption, mating success and predator evasion are discussed for each sex and related to fruit fly mating systems.     

Development time mediates the effect of larval diet on ageing and mating success of male antler flies in the wild

High-quality developmental environments often improve individual performance into adulthood, but allocating toward early life traits, such as growth, development rate and reproduction, may lead to trade-offs with late-life performance. It is, therefore, uncertain how a rich developmental environment will affect the ageing process (senescence), particularly in wild insects. To investigate the effects of early life environmental quality on insect life-history traits, including senescence, we reared larval antler flies (Protopiophila litigata) on four diets of varying nutrient concentration, then recorded survival and mating success of adult males released in the wild. Declining diet quality was associated with slower development, but had no effect on other life-history traits once development time was accounted for. Fast-developing males were larger and lived longer, but experienced more rapid senescence in survival and lower average mating rate compared to slow developers. Ultimately, larval diet, development time and body size did not predict lifetime mating success. Thus, a rich environment led to a mixture of apparent benefits and costs, mediated by development time. Our results indicate that ‘silver spoon'' effects can be complex and that development time mediates the response of adult life-history traits to early life environmental quality.     

Juvenile hormone regulation on the flight capability of Bactrocera dorsalis (Diptera: Tephritidae)

The oriental fruit fly, Bactrocera dorsalis (Hendel) (Diptera: Tephritidae), is considered a major economic threat in many regions worldwide. In order to better understand the flight capacity of B. dorsalis and its physiological basis, the functions and regulatory roles of juvenile hormone (JH) in the flight muscle of B. dorsalis were studied under a controlled environment. JH titer of B. dorsalis varied with age and sex. Females have higher JH titers and better flight capabilities than males, given that the increase in JH also corresponded to the ovarian development and maturation in females. The flight duration and distance of both males and females increased with the gradual increase of JH titer after adult emergences. Both JH titer and flight capability peaked in 15-d-old adult and declined subsequently with age. Flight activity stimulated the production of JH as adults flown for 24 h on the flight mills have the highest JH titers compare to adults flown on shorter flight durations. Adults treated with 0.5 µg and 5 µg of JH III were able to fly long durations and long distances, nevertheless when JH titer was too low or too high, it would restrict the flight ability of the fly. The mutual reinforcement of JH and flight activity provides fundamental understanding on the physiological aspects of the flight capability and dispersal, which facilitates strategies for the long-term control of this destructive pest.     

Effects of diet, ginger root oil, and elevation on the mating competitiveness of male Mediterranean fruit flies (Diptera: Tephritidae) from a mass-reared, genetic sexing strain in Guatemala

The release of sterile males is a key component of an areawide program to eradicate the Mediterranean fruit fly, Ceratitis capitata (Wiedemann), from Guatemala and southern Mexico. The objective of our study was to assess the effects of adult diet, exposure to ginger root oil (Zingiber officinale Roscoe), and elevation on the mating competitiveness of the sterile males used in an areawide program. Sterile males were maintained on a protein-sugar (protein-fed) or a sugar-only (protein-deprived) diet and were exposed (for 4 h 1 d before testing) or not exposed to ginger root oil. In field-cage trials conducted at a high (1,500 m) and low (700 m) site, we monitored the influence of these treatments on the mating success of sterile males in competition with wild males (reared exclusively on the protein-sugar diet and without ginger root oil exposure) for wild females. Elevation and ginger root oil exposure had significant effects, with sterile males having higher mating success at the low-elevation site and ginger root oil-exposed males having greater success than ginger root oil-deprived males at both sites. Diet did not have a significant overall effect, and its influence varied with elevation (dietary protein seemed to provide an advantage at the high-elevation site but not at the low-elevation site). Possible implications of these findings for eradication programs against the Mediterranean fruit fly are discussed.     

Role of the Corpus Allatum in the Control of Adult Diapause in the Blow Fly,Protophormia terraenovae

Adults of the blow fly, Protophormia terraenovae developed their ovaries promptly after emergence under long-day conditions at 25 degrees C, although they entered diapause under short-day conditions at 20 degrees C. Application of a juvenile hormone analog (JHA), implantation of the corpus allatum (CA) from reproductive adults, or transection of the nervus corporis allati induced ovarian development under diapause-inducing conditions. Removal of the CA suppressed ovarian development under diapause-averting conditions, and JHA application induced ovarian development in allatectomized adults. Therefore adult diapause in P. terraenovae is caused by a reduction of juvenile hormone secretion from the CA, and the endocrine activity of the CA is inhibited through a nervous pathway by the brain in diapause adults. Copyright 1997 Elsevier Science Ltd. All rights reserved     

取食重金属铜对棕尾别麻蝇亲代及子代生长发育与繁殖的影响

为了评估取食含重金属铜饲料对棕尾别麻蝇Boettcherisca peregrine亲代及子代的生长发育与繁殖的影响,在室内给棕尾别麻蝇初产幼虫饲喂含不同浓度（200, 400,800和1 600 µg/g）Cu²⁺的饲料直至化蛹,并对亲代和子代的生长发育和繁殖有关指标进行了观察和分析。结果表明：低浓度的Cu²⁺（200 µg/g）对其体重和体长起促进作用,但对幼虫历期、化蛹率、蛹历期、羽化率、性比、交配率和产仔量无显著作用;较高浓度的Cu²⁺ 则有抑制作用,且Cu²⁺处理浓度越高,亲代幼虫、蛹和雌雄成虫的体重越轻,幼虫和蛹的体长越短,化蛹率、羽化率、交配率和产仔量越低,幼虫历期和蛹期越长,成虫寿命越短。但Cu²⁺处理对成虫性比则无显著的影响。相比之下,经Cu²⁺处理后雌虫所产的子代若不再经Cu²⁺处理,其子代生存、生长发育与繁殖则基本不受影响,说明Cu²⁺对亲代的影响不能遗传至子代。此外,还探讨了该蝇亲代与子代体内Cu²⁺含量在其变态过程中的变化。     

The Effects of Temperature and Diet during Development,Adulthood, and Mating on Reproduction in the Red Flour Beetle

The effects of different temperatures and diets experienced during distinct life stages are not necessarily similar. The silver-spoon hypothesis predicts that developing under favorable conditions will always lead to better performing adults under all adult conditions. The environment-matching hypothesis suggests that a match between developmental and adult conditions will lead to the best performing adults. Similar to the latter hypothesis, the beneficial-acclimation hypothesis suggests that either developing or acclimating as adults to the test temperature will improve later performance under such temperature. We disentangled here between the effect of growth, adult, and mating conditions (temperature and diet) on reproduction in the red flour beetle (Tribolium castaneum), in reference to the reproduction success rate, the number of viable offspring produced, and the mean offspring mass 13 days after mating. The most influential stage affecting reproduction differed between the diet and temperature experiments: adult temperature vs. parental growth diet. Generally, a yeast-rich diet or warmer temperature improved reproduction, supporting the silver-spoon hypothesis. However, interactions between life stages made the results more complex, also fitting the environment-matching hypothesis. Warm growth temperature positively affected reproduction success, but only when adults were kept under the same warm temperature. When the parental growth and adult diets matched, the mean offspring mass was greater than in a mismatch between the two. Additionally, a match between warm adult temperature and warm offspring growth temperature led to the largest offspring mass. These findings support the environment-matching hypothesis. Our results provide evidence for all these hypotheses and demonstrate that parental effects and plasticity may be induced by temperature and diet.     

Male lures and male mating competitiveness in a genetic sexing strain of the melon fly (Diptera: Tephritidae)

The Sterile Insect Technique (SIT) has been used to suppress or eradicate populations of the melon fly, Zeugodacus cucurbitae (Coquillett), a pest of various fruits and vegetables throughout Asia, Africa, and South Pacific Islands. The success of the SIT depends largely on the ability of released, sterile males to compete successfully with wild males for copulations with wild females. Unfortunately, both the sterilization process (via gamma irradiation) and artificial selection under mass-rearing conditions generally reduce the biological competence, including the mating competitiveness, of the released males. However, previous data on Z. cucurbitae show that irradiation has minimal adverse effects and prolonged mass-rearing may or may not result in decreased mating ability. The present study first confirms that neither irradiation nor mass-rearing had deleterious effects on male mating ability of males from a pupal-color, genetic sexing strain (termed T1). Then, the influence of male lures (cue-lure and its natural analogue raspberry ketone) on the mating success of sterile T1 males was measured as a potential pre-release procedure to improve the effectiveness of SIT. Feeding on cue-lure enhanced male mating success, but the effect was short-lived (1 d but not 3 d post-feeding). Feeding on raspberry ketone boosted mating success over a longer time, i.e., at least 5 d but not 10 d post-feeding. However, mixing raspberry ketone into the adult diet had no detectable effect on male mating performance. The implications of these results for SIT against Z. cucurbitae are discussed.     

橘小实蝇交配行为观察

在实验室内对橘小实蝇Bactrocera dorsalis（Hendel）的交配行为进行了观察。描述了橘小实蝇交配中雄虫的“求偶场”,雄雄相遇、雌雌相遇、雌雄相遇时的相互反应,雌雄虫的交配行为与交配后行为。     

Total body nitrogen and total body carbon as indicators of body protein and body lipids in the melon fly Bactrocera cucurbitae: Effects of methoprene, a juvenile hormone analogue, and of diet supplementation with hydrolyzed yeast

The application of methoprene, and providing access to diet including hydrolyzed yeast, are treatments known to enhance mating success in the male melon fly Bactrocera cucurbitae Coquillett (Diptera: Tephritidae), supporting their use in mass rearing protocols for sterile males in the context of sterile insect technique (SIT) programmes. The objective of the present laboratory study was to investigate the effect of methoprene application and diet supplementation with hydrolyzed yeast (protein) on the turnover of body lipids and protein to confirm the feasibility of their application in melon fly SIT mass-rearing programmes. While females had access to a diet that included hydrolyzed yeast (protein), males were exposed to one of the following treatments: (1) topical application of methoprene and access to diet including protein (M+P+); (2) only diet including protein (M−P+); (3) only methoprene (M+P−) and (4) untreated, only sugar-fed, control males (M−P−). Total body carbon (TBC) and total body nitrogen (TBN) of flies were measured at regular intervals from emergence to 35 days of age for each of the different treatments. Nitrogen assimilation and turnover in the flies were measured using stable isotope (¹⁵N) dilution techniques. Hydrolyzed yeast incorporation into the diet significantly increased male body weight, TBC and TBN as compared to sugar-fed males. Females had significantly higher body weight, TBC and TBN as compared to all males. TBC and TBN showed age-dependent changes, increasing until the age of sexual maturity and decreasing afterwards in both sexes. Methoprene treatment did not significantly affect TBC or TBN. The progressive increase with age of TBC suggests that lipogenesis occurs in adult male B. cucurbitae, as is the case in other tephritids. Stable isotope dilution was shown to be an effective method for determining N uptake in B. cucurbitae. This technique was used to show that sugar-fed males rely solely on larval N reserves and that the N uptake rate in males with access to diet including hydrolyzed yeast was higher shortly after emergence and then stabilized. The implications of the results for SIT applications are discussed.     

Mating,insemination, and ovulation in the tsetse fly, Glossina morsitans

Virgin females of the tsetse fly, Glossina morsitans orientalis, retain their first egg within the right ovary whereas mated females ovulate. The component of the mating act which causes ovulation and thereby initiates the ovarian cycles which follow, is not insemination, the construction of a spermatophore in the uterus, or a humoral factor associated with the testes, accessory glands or ejaculatory ducts of donor males, or ‘full’ spermathecae of donor females. The ovulation rate increases with copulation time and females are shown to ‘add up’ their ‘sexual experience’ independently of that of the males. Inter-specific matings between G. morsitans and G. austeni also result in ovulation, but an unidentified factor associated with the completion of the mating act was also apparent. It is suggested that a ‘mechanical’ factor probably with nervous and endocrine components is responsible for the release of the egg from the ovary. Observations on the construction of empty spermatophores by aspermic males of G. morsitans are included.     

Diet alters species recognition in juvenile toads

Whether environmental effects during juvenile development can alter the ontogeny of adult mating behaviour remains largely unexplored. We evaluated the effect of diet on the early expression of conspecific recognition in spadefoot toads, Spea bombifrons. We found that juvenile toads display phonotaxis behaviour six weeks post-metamorphosis. However, preference for conspecifics versus heterospecifics emerged later and was diet dependent. Thus, the environment can affect the early development of species recognition in a way that might alter adult behaviour. Evaluating such effects is important for understanding variation in hybridization between species and the nature of species boundaries.     

Wheat germ oil and its effects on a liquid larval rearing diet for oriental fruit flies (Diptera: Tephritidae)

Wheat germ oil was added to a larval liquid diet for rearing Bactrocera dorsalis (Hendel) (Diptera: Tephritidae) to optimize fruit fly quality. Effects of various concentrations of wheat germ oil at 0.04, 0.07, 0.15, 0.30, and 0.66% and their possible mode of action were evaluated. Results suggest that addition of wheat germ oil does not affect pupal weight, larval developmental period, adult emergence, mating ability, or peak time for egg production. But there was a significant increase in pupal recovery, percentage of adult fliers, egg production, or egg hatch for larvae fed the diet with wheat germ oil compared with those reared on the liquid diet without wheat germ oil. The increase in egg hatch and fliers was dose dependent. Therefore, addition of wheat germ oil to fruit fly rearing diet is a novel way to improve fruit fly quality, especially in egg hatch, fliers, egg production, and pupal recovery.     

Methoprene application and diet protein supplementation to male melon fly,Bactrocera cucurbitae,modifies female remating behavior

Methoprene (an analogue of juvenile hormone) application and feeding on a protein diet is known to enhance male melon fly, Bactrocera cucurbitae Coquillett (Diptera: Tephritidae), mating success. In this study, we investigated the effect of these treatments on male B. cucurbitae's ability to inhibit female remating. While 14‐d‐old females were fed on protein diet, 6‐d‐old males were exposed to one of the following treatments: (i) topical application of methoprene and fed on a protein diet; (ii) no methoprene but fed on a protein diet; (iii) methoprene and sugar‐fed only; and (iv) sugar‐fed, 14‐d‐old males acted as controls. Treatments had no effect on a male's ability to depress the female remating receptivity in comparison to the control. Females mated with protein‐deprived males showed higher remating receptivity than females first mated with protein‐fed males. Methoprene and protein diet interaction had a positive effect on male mating success during the first and second mating of females. Significantly more females first mated with sugar‐fed males remated with protein‐fed males and females first mated with methoprene treated and protein‐fed males were more likely to remate with similarly treated males. Females mating latency (time to start mating) was significantly shorter with protein‐fed males, and mating duration was significantly longer with protein‐fed males compared with protein‐deprived males. These results are discussed in the context of methoprene and/or dietary protein as prerelease treatment of sterile males in area‐wide control of melon fly integrating the sterile insect technique (SIT).     

Separate and combined effects of nutrition during juvenile and sexual development on female life-history trajectories: the thrifty phenotype in a cockroach

We have yet to understand fully how conditions during different periods of development interact to influence life-history structure. Can the negative effects of poor juvenile nutrition be overcome by a good adult diet, or are life-history strategies set by early experience? Here, we tested the influence and interaction of different nutritional quality during juvenile and sexual development on female resource allocation physiology, life history and courtship behaviour in the cockroach, Nauphoeta cinerea. Nymphs were raised on either a good-quality or poor-quality diet. After adult eclosion, females were either switched to the opposite diet or remained on their original diet. We assessed mating behaviour and lifetime reproductive success for half of the females from each treatment. We evaluated reproductive investment, somatic investment and resource reallocation from reproduction to the soma via oocyte apoptosis in the remaining females. We found that poor juvenile conditions resulted in a fat phenotype with slow juvenile growth and short reproductive lifespan that could not be retrieved with a change in diet. Good juvenile conditions resulted in the converse, but again fixed, phenotype in adulthood. Thus, juvenile nutrition sets adult patterns of resource allocation.     

Structure,haemolymph titre,and regulatory effects of juvenile homone during ovarian maturation in Leucophaea maderae

Juvenile hormone (JH) synthesized and secreted in vitro by the corpora allata of mated adult Leucophaea maderae females was determined to be JH III (methyl-10,11-epoxy-3,7,11-trimethyl-2,6-dodecadienoate).The haemolymph titre of JH was determined during maturation of the terminal oöcytes in the first reproductive cycle of L. maderae. In virgin females, JH is not detectable in the haemolymph during the first eight days following adult emergence; however, by 10 days after emergence, trace quantities of JH are apparent. Mating stimuli induce a dramatic increase in the concentration of haemolymph JH, with a peak occurring approximately 12 days after mating; thereafter, the JH concentration declines until it has reached an undetectable level 19 days after mating, at the time of chorion deposition.During ovarian maturation, changes in the rates of synthesis of vitellogenin by the fat body and DNA by the ovary correlate closely with the haemolymph titre of JH. However, no such correlation exists between the JH titre and the extensive ovarian protein synthesis that occurs in L. maderae coincident with chorion formation.The effects of JH I and JH III on both vitellogenin synthesis and ovarain DNA synthesis are statistically similar.     

Drosophila americana Diapausing Females Show Features Typical of Young Flies

Diapause is a period of arrested development which is controlled physiologically, preprogrammed environmentally and characterized by metabolic depression that can occur during any stage of insect development. Nevertheless, in the genus Drosophila, diapause is almost always associated with the cessation of ovarian development and reproductive activity in adult females. In this work, we show that, in D. americana (a temperate species of the virilis group), diapause is a genetically determined delay in ovarian development that is triggered by temperature and/or photoperiod. Moreover, we show that in this species diapause incidence increases with latitude, ranging from 13% in the southernmost to 91% in the northernmost range of the distribution. When exposed to diapause inducing conditions, both diapausing and non-diapausing females show a 10% increase in lifespan, that is further increased by 18.6% in diapausing females, although senescence is far from being negligible. ActinD1 expression levels suggest that diapausing females are biologically much younger than their chronological age, and that the fly as a whole, rather than the ovarian development alone, which is phenotypically more evident, is delayed by diapause. Therefore, diapause candidate genes that show expression levels that are compatible with flies younger than their chronological age may not necessarily play a role in reproductive diapause and in adaptation to seasonally varying environmental conditions.