Nectar-related vs. human-related volatiles: behavioural response and choice by female and male Anopheles gambiae (Diptera: Culicidae) between emergence and first feeding

The close association of Anopheles gambiae Giles with humans and its females' ability to live on human blood alone suggest that females may ignore sources of sugar in favour of human blood as a source of energy. They have limited energy reserves at emergence, and at 27 degrees C both sexes generally die if they do not feed during night 1, 24-36 h after emergence. Food preferences during this critical period were tested by measuring responses to volatiles from honey and soiled socks, which served as surrogates for nectar-related and human-related volatiles in a wind-tunnel olfactometer. Both sexes responded more strongly to honey than to human volatiles, and given a choice, preferred honey over human volatiles. After 5 days of sugar access and maturation, males continued to prefer honey volatiles, whereas females changed behaviour, responding almost exclusively to human volatiles. Night 1 experiments also demonstrated that: (i). females previously having had sugar during the night of emergence responded more strongly to human volatiles; (ii). large-bodied mosquitoes of both sexes responded more strongly to honey than small-bodied ones; and (iii). females were equally responsive to honey in both early and late scotophase but were slightly more responsive to human volatiles in late scotophase. These results indicate that for a female's first meal, sugar is a viable option and is preferred when nectar-related stimuli are strong. This supports field evidence that sugar-feeding is a significant component of A. gambiae female behaviour.     

Reproductive physiology of Anopheles gambiae and Anopheles atroparvus.

When exposed to a human host, Anopheles gambiae started probing 4 h post-eclosion, but 95% successfully blood-fed by 16-20 h with maximal blood volumes of 5- 10 microl per female. When fed sugar, the 95% feeding was not observed until 36-40 h post-eclosion; sugar meals appeared to interfere with blood meals. Similarly in An. atroparvus, maximum volumes were 10 microl when starved but only 6 microl when fed sugar. This species did not bite before 2 d, and 95% biting was by 4 d. Given single blood meals to water-kept An. gambiae, a threshold body size for oogenesis was detected. With wing lengths below 2.8 mm, eggs never matured, but when sugar-fed, females of all sizes matured eggs including the synthesis of maternal deposits. Although sugar feeding interfered with blood feeding, more lipid was transferred to the yolk. In water-kept An. atroparvus only 5% of the females produced eggs. When sugar-fed for 4 d, all females matured eggs, so in this species sugar feeding appeared to be essential for oogenesis. An. gambiae always took multiple blood meals, tested at any time after the first ones, leading to 120 mature eggs/female. Yolk composition was 3.9 mcal protein and 3.8 mcal lipid/oocyte when kept on water, but 2.8 meal protein and 4.3 mcal lipid/oocyte with intermittent sugar meals, thus marking a surprising flexibility in synthesis of yolk protein and lipid that strongly depends on additional carbohydrates sources. Only 80% of water-fed An. atroparvus re-fed 2 d after a first blood meal with small females taking three blood meals but they still showed reduced fecundity. Only the large water-fed females matured eggs, with blood volumes higher than 9-12 microl. When fed sugar, the blood meal input was reduced, but oogenesis was possible, whereas water-fed females required three blood meals to reach the caloric level comparable to pre-feeding sugar-fed females. Water-fedAn. gambiae could survive on daily blood meals alone, but survival was further extended by intermittent sugar meals. When offered a blood donor daily, there was a behavioral difference. Females maintained alone showed a more or less regular 3 d feeding and oviposition activity, while females kept in groups fed daily followed a daily oviposition pattern, suggesting gonotrophic discordance.     

A survival and reproduction trade-off is resolved in accordance with resource availability by virgin female mosquitoes

The first 2-4 days after an Anopheles gambiae female mosquito emerges are critical to her survival and reproductive success. Yet, the order of behavioural events (mating, sugar feeding, blood feeding) during this time has received little attention. We discovered that among female cohorts sampled from emergence, sugar feeding had a higher probability than blood feeding of occurring first, and mating rarely occurred before a meal was taken. The night after emergence, 48% of females fed on sugar in mesocosms, and 25% fed on human blood; in the absence of sugar, 49% of females fed on human blood. After 5 days, 39% of the sugar-supplied females had blood fed and mated, and were fructose negative, whereas only 8% of the sugar-denied females had both blood fed and mated by this time. The model that best explained the transitions suggests that females made use of two distinct behavioural pathways, the most common one being to sugar-feed, then mate, and then seek blood. Other females sought blood first, then mated, and forwent a sugar meal. Lipid levels were higher in females with access to sugar than in females without access to sugar, particularly for those in later gonotrophic stages, while glycogen levels in the sugar-supplied group were higher throughout. In single-night experiments with females having had access to sucrose since emergence, those given a blood meal 1 day before spending a night with males had higher insemination rates than those not receiving the blood meal. These results indicate that the trade-off between survival and immediate reproduction is resolved by young adult females in accordance with availability of resources and gonotrophic state.     

Lipid metabolism during sequential gonotrophic cycles in large and small female Aedes aegypti

The lipid metabolism was investigated during six gonotrophic cycles of Aedes aegypti. Females of constant body size were analyzed for their total lipid content: large females with a body size of 41.06 (wing length cubed) and small females with 15.63. Their lipid contents at eclosion were compared to lipid values after two days of sugar-feeding, shortly before a blood meal, after oviposition, of their total egg batches, and again before the next blood meal, with intermittent access to sugar for two days for six gonotrophic cycles.Large females transferred most of their pre-blood meal lipid into the ovaries. Their low lipid content after oviposition was restored by synthesis from intermittent sugar meals. After the third gonotrophic cycle, they withheld more and more of the resynthesized lipid in their fat body, thus gradually reducing their fecundity. Since blood consumption was not altered significantly during these six cycles, age-related reduction of fecundity was clearly caused by limitations of yolk lipid.Small females transferred a considerably smaller, but constant segment of sugar-derived lipids to the ovaries. In both size classes, lipid content per oocyte was constant throughout all cycles with 9 mcal/oocyte in large and 7 mcal/oocyte in small females. Total fecundity reached 450 eggs in large and 280 eggs in small females. Large females that were maintained on water without sucrose took large blood meals from which part of the yolk lipid was synthesized. Extrapolations suggest that only one or two additional gonotrophic cycles would be possible without additional carbohydrate sources, despite lipogenesis from blood protein.     

Anopheles gambiae feeding and survival on honeydew and extra-floral nectar of peridomestic plants

It is widely believed that the malaria vector Anopheles gambiae Giles (Diptera: Culicidae) rarely or never feeds on sugar in nature. If so, the need for supplemental blood-feeding may be increased and this would help to explain why it is such an efficient malaria vector. Nonetheless, both sexes of this mosquito species readily imbibe and digest sugar solutions, and sugar is a staple of laboratory colonies. In this study, we investigated whether An. gambiae will feed on the extra-floral nectar of three common peridomestic plants in Africa, and on honeydew of the mealybug Pseudococcus longispinus (Targioni-Tozetti) (Hemiptera: Homoptera: Pseudococcidae), and how this affects survivorship. We found that both males and females of An. gambiae provided with vegetative parts of cassava (Manihot esculenta Crantz) survived as well (x = 26.3 and 19.2 days, respectively) as they did on 50% sucrose solution (x = 29.7 and 24.3 days, respectively) and much longer than they did on water alone (x = 1.8 days, both sexes). Females provided with mealybug honeydew also lived substantially longer (x = 16.5 days) than those on water alone. Males and females provided with vegetative parts of castorbean (Ricinus communis L.) also survived much longer (x = 12.7 and 7.8 days, respectively) than on water, but those provided with flowering lantana (Lantana camara L.) did not. Anthrone tests of females after one night of exposure to these potential energy sources confirmed that they obtained fructose from cassava, from mealybug honeydew, and from non-flowering castorbean, but not from lantana or from castorbean lacking its petiolar nectaries. Previous laboratory studies had shown that sugar availability affects the survival and biting frequency of An. gambiae. It now appears that this mosquito can locate natural sources of plant sugar readily and utilize them effectively. Nectar-producing plants in the domestic environment may play a significant role in this mosquito's energy budget and malaria vectorial capacity.     

Effect of body size and sugar meals on oviposition of the yellow fever mosquito,Aedes aegypti (Diptera: Culicidae)

The effects of dietary sugar and body size on the oviposition of Ae. aegypti were studied under laboratory conditions. In female mosquitoes provided with sugar, the start of maximum fecundity was significantly delayed and the oviposition period was longer than in females provided with water. The peak of oviposition was also delayed in sugar‐fed females. Large females oviposited more eggs per day than small females at maximum fecundity and during eight days of observations. Large females also visited significantly more water‐containing cups in their cages per day than small females at maximum fecundity. During the eight days of observations, large females and sugar‐fed females visited more water‐containing cups in their cages than water‐fed small females. Both large females and sugar‐fed females oviposited their eggs at sites higher above the water line than water‐fed small females. These results suggested that large and sugar‐fed female Ae. aegypti mosquitoes had more energy reserves and oviposited their eggs at higher sites, which would lead to a time lag in hatching.     

Life history and demography of the predatory mite,Amblyseius longispinosus Evans

The life history and demography of Amblyseius longispinosus Evans were studied under laboratory conditions 25–28°C and 65–85% RH using the red form of the twospotted spider mite, Tetranychus urticae Koch complex as prey. The entire development from egg to adult averaged 102.5 h for both sexes with a survival rate of 90.0%. Immature females consumed more prey eggs than males, averaging 7.9 and 6.4 eggs/day, respectively. The embryonic development time for male eggs was longer (average 45.2 h) than for female eggs (average 42.6 h). The average duration of succeeding stages did not differ between the sexes: larva, 15.7; protonymph, 21.1; and deutonymph, 23.0 h. Larvae were not observed to feed. Statistically significant differences between sexes were noted with respect to consumption of the deutonymphs (female, average 4.0±0.2; male, average 2.9±0.3 eggs/day) but not of the protonymphs. Mating occurred on the same day as adult emergence and was repeated several times during the reproductive life. Egg laying started on the second day after emergence. Parameters relating to oviposition were: fecundity, gross (50.7) and net (43.3) eggs/female; hatchability, 99.6% (gross) and 99.5% (net); the oviposition lasted at most 28 days and showed a peak in the first week. Net reproductive rate (R₀) was 36.7 female offspring/female/generation time (T) of 9.0 days. The sex ratio was biased toward the females (71.9%), while the intrinsic rate of increase (r_m) was 0.4 with a doubling time (DT) of 1.7 days. Males lived longer (36 days) than females (30 days) with a LT₅₀ for males of 26 days and for females of 22 days.     

Host-feeding and its effects on the physiology and behaviour of the ichneumonid parasitoid, Coccygomimus turionellae

ABSTRACT. Coccygomimus turionellae was found to possess a pre-ovipositional period of 3–6 days during which it ignored hosts. This period corresponded to the time needed for mature eggs to develop. Once oviposition began, the females exhibited host-feeding behaviour, using the nutrients for oogenesis. Host-deprived females were able to mobilize some body tissue in the production of eggs, but this resulted first in the loss of their flight ability and then in an increasing paralysis of their hind legs. Host-feeding varied from the females only sipping at the wound up to the female ramming her ovipositor around in the wound and/or chewing at the drill site. Wasps that only fed at the wound usually deposited eggs and those that had a high rate of host contact used the hosts mainly for oviposition. Wasps that had been without hosts showed a much greater tendency to feed, often destroying the hosts. The greatest preference for feeding was on hosts that were also highly preferred for oviposition. However, when less desirable beetle pupae were accepted, they were fed on to a larger degree than were the more ovipositionally desirable moth pupae. Host deprived parasitoids demonstrated an increased tendency to feed on hosts that are never naturally used for oviposition.     

Influence of feeding interval on reproduction and longevity ofPodisus sagitta (Het.: Pentatomidae)

The effect of various feeding intervals on reproduction and longevity of the predatory bugPodisus sagitta (Fabricius) (Heteroptera: Pentatomidae) was studied in the laboratory. Isolated pairs of adult bugs were given moisture in excess and were fed 1 seventh-instar larva of the greater wax moth,Galleria mellonella (L.), with a mean weight of 90 mg every 1, 3, 7, 14 and 21 days. Reproduction ofP. sagitta reflected well the feeding intervals. Females fed at increasing time intervals reduced reproductive outputs by decreasing oviposition frequency and size of egg batches. Predators fed every 21 days produced a mean of only 1.0 egg per day in comparison with 9.9 eggs per day for females fed daily. Longevity of females fed daily was 110.7 days; females fed less frequently tended to live longer, with a maximum mean longevity of 217.5 days for females fed every 14 days. No cannibalism among adults was observed, but females were occasionally seen to feed on their own eggs. Availability of moisture was found to be essential for the predators to survive during periods of low food supplies. These findings are discussed in relation to the predator's adaptation strategies in conditions of prey scarcity.      

Biology and reproductive parameters of the brown lygodium moth, Neomusotima conspurcatalis--a new biological control agent of Old World climbing fern in Florida

Neomusotima conspurcatalis Warren was first released in Florida as a weed biological control agent against Old World climbing fern in 2008, and readily established large field populations. A related biocontrol agent, Austromusotima camptozonale, had previously failed to establish despite several years of releases. Life history studies were conducted to determine whether aspects of the reproductive biology of N. conspurcatalis might account for these different outcomes. At 26.5°C, development from egg to adult averaged 22.2 ± 0.1 d, with 75% of larvae emerging as adults. The sex ratio averaged 1:0.8 (♂:♀), with both sexes emerging at the same time. Female moths typically mated once, on the first night after emergence, and began oviposition the next night. Females laid half their eggs on the first night and lived an average of 10.7 ± 0.8 d. Individual females maintained in cages with a male-biased sex ratio (3♂:1♀) produced significantly more larvae over their lifetime (140 ± 6.6 larvae) than individual females maintained at a ratio of 1♂:1♀ (111 ± 9.1 larvae). Sexual selection, either through 'male-male competition' or 'female choice' was likely responsible for this result, because there were no significant differences in mating frequency, duration of ovipositional period or female longevity to otherwise explain the difference. Two-fold greater lifetime reproductive output (average 127 ± 6.3 larvae) and deposition of half this output on the first night of oviposition, likely contributed to rapid field establishment of N. conspurcatalis compared with A. camptozonale.     

Field observations of mating,oviposition, and feeding behavior ofAnticarsia gemmatalis (Lepidoptera: Noctuidae) in a soybean field

The temporal occurrence and frequency of mating, oviposition, and feeding of wild velvetbean caterpillar moths, Anticarsia gemmatalis,in a soybean field (Glycine max)were documented. Mating occurred predominately during the first 4 h after sunset and primarily on leaflet bottoms. Oviposition was concentrated within the first 6 hr after sunset. Eggs were laid singly, and in areas of high trichome density, on leaflets, pulvini, and petioles. Feeding occurred primarily at night, with most males feeding during the first half of the night and females during the second half. Males and females shared some food sites but others were visited strictly by males.     

Effects of age and mated status on flight potential of Helicoverpa armigera (Lepidoptera: Noctuidae)

Abstract The effects of ageing and female mated status on the flight potential of Helicoverpa armigera (Hübner), collected as larvae from a pigeonpea crop in southern India, were investigated using a tethered-flight technique. In non-mated moths fed sugar solution, from the first night after adult eclosion, the durations of both total and longest continuous flight per night increased up to night 4 and remained at this level until at least night 6. Ovarian maturation was rapid with 77% of unmated moths having commenced oviposition by the third night. On the basis of field evidence it is likely that most females would be mated by the third or fourth night, provided plants with nectar or sugary exudates were locally available. In successfully-mated females a 15-fold decrease in total flight duration and a 28-fold decrease in longest continuous flight duration was observed in contrast to non-mated females of similar age. As host plants suitable for adult feeding and oviposition were locally available during the time of feral adult emergence, synchronous pre-reproductive migration was unlikely to occur in the population studied.     

Does the Anopheles blood meal-fecundity curve, curve?

We determined the blood meal size-fecundity relationship in Anopheles gambiae in the laboratory. Our primary interest was to determine whether the fecundity curve has a non-linear component, i.e.. does it decelerate towards an asymptote? Small and large adult females in their second gonotrophic cycle were fed on human hosts for predetermined lengths of time. Blood meal size was ascertained by weight and subsequent fecundity was determined by the number of eggs laid on filter paper. In both small and large-bodied females there was a significant curvilinear component. We used this relationship to show that marginal returns from feeding decline as a function of current blood load. This means that further fecundity payoff for continued feeding declines over time.     

草地贪夜蛾对水稻和玉米的取食和产卵选择性与适合度

【目的】明确草地贪夜蛾Spodoptera frugiperda对寄主植物水稻和玉米的取食和产卵选择性与适合度,进而分析草地贪夜蛾对水稻的为害风险。【方法】采用自由选择法,测定草地贪夜蛾对水稻和玉米的取食、产卵选择性。调查取食水稻和玉米苗的草地贪夜蛾的生长发育、存活率、繁殖力的差异,用种群增长趋势指数(I)评价水稻对草地贪夜蛾的适合度。【结果】草地贪夜蛾对水稻和玉米的取食和产卵选择性存在显著差异。接虫后2-48 h,幼虫对玉米的取食选择率随时间延长而逐渐增加,而幼虫对水稻的取食选择率随时间延长而逐渐降低;接虫后48 h,2龄幼虫和4龄幼虫对玉米的取食选择率分别为93.06%和59.72%,分别是对水稻的取食选择率的66.95倍和21.48倍。自由产卵6 d,在玉米上的产卵量是在水稻上的8.64倍。与取食玉米苗的相比,取食水稻苗的草地贪夜蛾幼虫发育历期延长,卵孵化率、幼虫存活率、化蛹率和成虫羽化率均显著降低,雌性比率下降,雌成虫寿命显著缩短,单雌产卵量显著减少。取食玉米苗的草地贪夜蛾的种群增长趋势指数(I)为165.93,其在水稻上的相对适合度为0.21。【结论】相较于水稻,草地贪夜蛾在玉米上表现出更高的适合度,其偏好在玉米上取食和产卵。水稻不是目前侵入我国的草地贪夜蛾的嗜食寄主,但其可以通过取食水稻幼苗正常生长发育并完成生活史,在其种群密度较大、嗜食寄主植物缺乏时存在转移为害水稻的潜在风险。     

Feeding and reproduction of XX Ctenophthalus wagneri Tifl., 1928 (Ctenophthalmidae, Siphonaptera) in experiments]

Under laboratory conditions at 20 to 21 degrees C young imagos of C. wagneri proceed quickly to feeding and reproduction. They suck blood about three times a day and females oviposit three times, each laying consisting of three eggs. When placed in a refrigirator at 2 to 5 degrees some young individuals lay eggs, while others do not start oviposition up to 5 months (observation period). Fleas, which already laid eggs under laboratory conditions, being transferred into a refrigirator decrease the degree of their attachment to the host and feeding frequency (up to one time a day). Oogenesis of females is inhibited and most of them cease egg laying.     

The nutritional contribution of males affects the feeding behavior and spatial distribution of females in a bruchid beetle,<Emphasis Type="Italic"> Bruchidius dorsalis</Emphasis>

The relationship between the quality/quantity of male investment and the feeding behavior of females was investigated in a bruchid weevil, Bruchidius dorsalis (Fahraeus), whose males donate nutrition via seminal fluid to females. Experiments on the effect of feeding regimes of both sexes on the mating frequency of females showed that females mated at a higher frequency if given low-quality food or poor male investment. On the other hand, the experiment that examined the effect of male investment quality on female feeding behavior showed that females receiving the high-quality investment exhibited feeding behavior less often. These results suggest that male investment and feeding behavior play the same role for B. dorsalis females. These experiments also showed that there are sex-related asymmetries in mating and feeding behaviors: females mated more often but males fed more often. Moreover, a field census suggested that only males visited non-host flowers to feed on the pollen and nectar during the non-flowering period of the host plants; females always stayed on the host plants irrespective of the flowering phenology. These results suggest that in B. dorsalis courtship role reversal and sex-specific feeding modes are fairly fixed and obligatory, and that male investment, derived from sexual selection, could affect the feeding behavior and spatial distribution of both sexes, which may have far-reaching impact in various ecological contexts.     

Feeding behavior and reproduction of Stenophthalmus wagneri Tifl., 1928 (Stenophthalmidae, Siphonaptera) under experimental conditions]

Under laboratory conditions at 20 to 21 degrees C young imagos of C. wagneri proceed quickly to feeding and reproduction. They suck blood about three times a day and females oviposit three times, each laying consisting of three eggs. When placed in a refrigirator at 2 to 5 degrees some young individuals lay eggs, while others do not start oviposition up to 5 months (observation period). Fleas, which already laid eggs under laboratory conditions, being transferred into a refrigerator decrease the degree of their attachment to the host and feeding frequency (up to one time a day). Oogenesis of females is inhibited and most of them cease egg laying.     

Faeces feeding by adult Phormia regina (Diptera: Calliphoridae): impact on reproduction

resh beef liver, sugar, and five different types of faeces were evaluated as supportive diets for egg development in the blowfly Phormia regina . Females on a sugar diet were unable to develop follicles beyond stage 3, whereas liver proved to be the best diet for complete egg maturation. Some faecal diets were unable to support egg maturation when fed upon for a short period of time; however, longer periods of feeding produced complete egg maturation. The necessity to feed for longer periods of time in order to produce eggs on most of these diets was attributed to their low protein content. Males, in a shorter period of time than females, obtained enough protein from faeces to activate the neuroendocrine system involved in mating.     

Effect of bacteria on nutritional status and reproductive success of the Mediterranean fruit fly Ceratitis capitata

Abstract.  Mediterranean fruit flies ( Ceratitis capitata Wiedemann, Diptera: Tephritidae) harbor a community of diazotrophic bacteria in their digestive system. The present study aims to test the hypothesis that bacteria contribute to fly fitness by enhancing copulatory success and egg production in males and females, respectively. After eclosion, flies were fed a full diet containing peptides, sugar and minerals, or a sugar diet, lacking peptides. Subgroups from each diet were fed a mixture of the antibiotics ciprofloxacin and piperacillin. The presence of bacteria, food consumption, weight gain, lipid and protein levels, oviposition in females and copulatory success of males were quantified in the four groups. The antibiotic treatment effectively cleared the gut of bacteria. The relative amounts of food consumed (with or without antibiotics) are similar in all groups. The antibiotics do not inhibit feeding, and their ingestion does not affect dry weight or the amount of protein stored, yet females feeding on the full diet without antibiotics have increased lipid levels. Females fed the full diet produce significantly more eggs than females on the sugar diet, but the presence of bacteria does not affect numbers of eggs produced. However, in the absence of bacteria, the oviposition rate of nutritionally stressed females is significantly accelerated. The presence of bacteria in sugar fed males does not provide them with a mating advantage. Conversely, in males fed a full diet, the presence of bacteria is associated significantly with a shorter latency to mate. It is concluded that, because the bacterial community is present at all stages of the fly's life cycle, at least some species are effectively transmitted from parents to offspring, and removal of bacteria affects measurable physiological and behavioural parameters related to fitness, the association between bacteria and the medfly is mutualistic.