Induction of adult diapause and of low and high reproductive states in a parasitoid wasp,Ooencyrtus nezarae, by photoperiod and temperature

Ooencyrtus nezarae Ishii (Hymenoptera: Encyrtidae) was reared on eggs ofRiptortus clavatus (Thunberg) (Heteroptera: Alydidae) at various temperatures under long-day (L16:D8) or short-day (L10:D14) conditions. There was no diapause during egg, larval or pupal stages under any set of conditions examined. However, at 15°C under short-day conditions, vitellogenesis was arrested in all adult females and they entered diapause. At 15°C under long-day conditions, or at 20°C under short-day conditions, some adult females entered diapause. Under the latter set of conditions, the adult females laid eggs but they laid fewer eggs than under long-day conditions, Even at 25°C, under short-day conditions, adult females laid fewer eggs than under long-day conditions, and this low rate of oviposition was attributed to the retarded development of ovaries. Diapause adults reared at 15°C were more resistant to low temperature than nondiapause adults reared at 25°C.     

Photoperiod and temperature studies to determine whether diapause is found in successive generations of the African phytoseiid,Euseius fustis (Pritchard and Baker) (Acari: Phytoseiidae)

Photoperiod and temperature conditions known to induce diapause in tropical arthropods were tested on two generations (G₀ and G₁) of the phytoseiid, Euseius fustis. Failure to lay eggs or a pre-oviposition period longer than 15 days were the criteria used to determine whether females were in diapause. Females reared from egg to adult and held throughout adult life under cyclic temperatures of 29/20°C in combination with long photophases of 16L: 8D and 14L: 10D showed no indication of aestival diapause. Similarly, hibernal diapause was not induced in females reared under a constant temperature of 18°C and a photophase of 8L: 16D. Under the various test conditions, females initiated oviposition within an average of 4 days. Overall, pre-oviposition patterns for G₀ and G₁ females were similar under the same test conditions. Reproductive patterns based on the mean number of eggs per female per day varied only slightly between generations for the same treatments. No behavioural or morphological attributes associated with diapause were observed.     

The induction of diapause in <Emphasis Type="Italic">Moina</Emphasis> by species-specific chemical cues

The ability to change the reproduction mode and produce diapausing eggs, which is prevalent in many zooplankton species, significantly impacts on the evolution and ecology of aquatic communities. The production of diapausing eggs is controlled by multiple effects of biotic and abiotic factors, including infochemicals. We have investigated the effects of chemicals exuded by conspecifics and ecologically close competing congers, Moina brachiata and M. macrocopa, which coexist in the same water body, and by larger Cladocera species (Daphnia magna) on the change of reproduction mode, specific growth rate and fecundity of M. brachiata and M. macrocopa females. The production of gametogenetic eggs in both species was detected only in waters from crowded cultures of conspecifics. The water from crowded cultures of conspecifics reduced the specific growth rate of the juvenile females of both species that later switched to gametogenesis. While it either did not affect (in M. macrocopa) or even increase (in M. brachiata) the specific growth rate of the juvenile females that later reproduced by parthenogenesis. Females of M. macrocopa released significantly fewer neonates in the water from crowded cultures of conspecifics than in all other treatments, while the fecundity of M. brachiata females was the same in all treatments. To understand the phenomenon of diapause induction under the effect of chemical cues in zooplankton, a link between laboratory tests and ecological research should be established, and the chemical composition of the signals should be determined.     

Effect of relative humidity on development, survival and reproduction of the predatory lacewing Dichochrysa prasina (Neuroptera: Chrysopidae)

The effect of five constant relative humidity (RH) regimes (12%, 33%, 55%, 75% and 94%) on the preimaginal development and adult longevity and reproduction of Dichochrysa prasina Burmeister was studied at 26 ± 1 °C and a photoperiod of 16:8 (L:D) under laboratory conditions. For the experiments, larvae were fed individually on eggs of the flour moth Ephestia kuehniella (Zeller), whereas adults on a liquid diet consisting of water, yeast hydrolysate, sugar and honey. The experimental insects were maintained individually in sealed plastic containers with saturated water salt solutions at the base to control RH. Preimaginal developmental time from egg to adult for females and males was significantly shorter at 75% and 94% RHs than at other RHs in the range from 12% to 55%. Percentages of adult emergence were similar at all RHs tested and varied from 52.5% to 75.0%. Adult females that had no access to water and were maintained at 12%–75% RH lived on average less than two weeks and laid no eggs, whereas at 94% RH they lived longer and each laid on average 201 eggs. When the females had periodic access to a water source a significant increase in both longevity and egg production occurred and the estimated intrinsic rate of increase was high, irrespective of RH. These results could be useful for the mass-rearing of D. prasina and for better understanding its population dynamics under field conditions.     

Maternal effects, conspecific chemical cues, and switching from parthenogenesis to gametogenesis in the cladoceran Moina macrocopa

The change in reproductive mode from parthenogenesis to gametogenesis in Cladocera is controlled by multiple environmental cues. Maternal effects are involved in the control of reproductive switching. In this study, we estimated the readiness of Moina macrocopa females to change reproduction mode under the effect of conspecific chemicals on maternal and offspring generations. The results demonstrated that information about the chemical environment was not transmitted between generations (none of the females produced diapausing eggs in the control medium irrespective of their mothers' environment). Differences in maternal energetic investments were not significant, hence the maternal environment did not affect the fecundity. However, tested animals adapted to the effect of the diapause inducing factor. When offspring of mothers cultured in crowded water were also cultured in crowded water (the constant effect of conspecific chemicals), they switched less readily to gametogenesis than offsprings of mothers cultured in fresh medium.     

Seasonal Variation in Diel Production Pattern of Centropages tenuiremis in Xiamen Waters, China

The seasonal variations in diel production pattern, and egg hatching time of the copepod Centropages tenuiremis in Xiamen waters, China, were investigated between January and June of 2003. The results show that in the winter–spring (January to early May) the adult females tended to lay subitaneous eggs at night, while in the beginning of summer (late May–June) they tended to lay diapause eggs as well as subitaneous eggs during the daytime. The egg hatching time negatively correlated with water temperature. These results, combined with the fact that C. tenuiremis migrates vertically in Xiamen waters, may explain the reproductive strategy of this dominant species in winter–spring.     

The production of subitaneous and diapause eggs: a reproductive strategy for Acartia bifilosa (Copepods: Calanoida) in Southampton Water, UK

Acartia bifilosa from Southampton Water lays two morphologically distinct types of egg which are described for the first time. Eggs with a smooth surface are considered subitaneous, while eggs covered with thick 'spines' are diapause. During the seasonal occurrence of A.bifilosa in Southampton Water, from November/December to June, subitaneous eggs are laid during the first months of this period. The production of diapause eggs is restricted to a 2 month period before A.bifilosa disappears from the water column. There are significant differences between the response of the eggs produced at seasonal field temperatures (5-12C) and those reported for other Acartia species. In particular, the numbers of eggs female^-1 day^-1, both subitaneous and diapause, are lower and typically <4; and the hatching time of subitaneous eggs is longer, at up to 10 days, at optimum temperatures between 15 and 20°C and optimum salinity >20 PSU. Females acclimated to higher laboratory temperature regimes show higher egg production rates at field salinity. A delayed-hatch subitaneous egg is also reported. Results suggest that fecundity in A.bifilosa from Southampton Water might be limited to some degree by temperature and the responses of the eggs produced appear to offer A.bifilosa a reproductive repertoire to ensure its sustained presence in this seasonally influenced environment.      

Oviposition behavior of Edovum puttleri, reared on two hosts, Leptinotarsa decemlineata and L. texana

Female Edovum puttleri Grissell [Hymenoptera: Eulophidae], reared from eggs of Leptinotarsa decemlineata (Say) or Leptinotarsa texana Schaeffer [Coleoptera: Chrysomelidae], were videotaped as they attacked egg masses of L. decemlineata containing 20 host eggs. We identified 15 components of ovipositional behavior. Parasitoids reared on L. texana attacked and oviposited in significantly more host eggs than did females reared on L. decemlineata. Ethometric analyses of behavioral transitions and a clustering analysis of 34 behavioral parameters showed that females reared on L. texana attacked the host egg mass in a different manner than those reared from L. decemlineata. It was concluded that differences were associated with the host species upon which they were reared. Contrary to previous reports, mortality of unparasitized hosts was caused by an ovipositor probe of short duration, which was not related to host-feeding.     

Is juvenile hormone involved in the maternal regulation of egg size and progeny characteristics in the desert locust?

The role of juvenile hormone (JH) in the maternal regulation of progeny characteristics was examined in the desert locust, Schistocerca gregaria. Female adults of this species are known to produce smaller but more eggs when reared in isolation than do those reared in a group. Eggs laid by isolated females develop green hatchlings typical of solitarious forms, whereas those laid by the latter produce black hatchlings typical of gregarious forms. Topical application of a juvenile hormone analog (JHA), fenoxycarb, or implantation of corpora allata (CA) taken from the migratory locust, Locusta migratoria, caused crowded S. gregaria females to deposit smaller eggs, but did not have a significant effect on the number of eggs per egg pod except at high doses of JHA. The production of smaller eggs by treated and untreated crowded females was closely associated with earlier deposition of the egg pods and shorter oviposition intervals. However, neither JHA application nor CA implantation influenced the progeny characteristics in actively reproducing aged females under crowded conditions, while untreated control females started producing smaller and more eggs upon transfer to isolated conditions. These results may suggest that JH is not directly involved in the maternal regulation of phase-dependent progeny characteristics.     

Mating and fecundity of Dermatophagoides farinae

Studies of the life cycle of cultured Dermatophagoides farinae found that after an initial mating D. farinae females lived for 63.3 ± 64.6 (SD) dafter their egg production period ended .The long period after cessation of egg production for D. farinae suggested D. farinae females could mate multiple times and produce eggs continuously for a longer period. The purpose of this study was to determine if female D. farinae could mate at least two times, and subsequently increase the production of viable eggs over a longer period of time compared to a single mating. Female D. farinae were randomly selected from thriving cultures and isolated in cages. When the females had ceased to lay eggs a male was added to the cage. Fifty-seven percent of the isolated females mated again and produced a second batch of viable eggs. In natural or culture populations, females have continuous availability of males. Therefore, in another experiment, females that emerged from the tritonymphal stages were continuously exposed to fresh males and fecundity and lengths of the reproductive and post reproductive periods were determined. These females had a 11 d longer reproductive period and produced 30.7% more eggs compared to females that only mated one time after they emerged from the tritonymphal stage. However, the post reproductive period was still long (58.6 ± 11.4 [SE] d) the significance of which is not clear. In conclusion, this study revealed that D. farinae females are capable of more than one successful mating that results in increased egg production compared to that of a single mating. It is likely that females mate multiple times in natural and culture populations. It was observed that females actively attract males during the reproductive period but not afterward even though she continues to live a long time. This revised version was published online in July 2006 with corrections to the Cover Date.     

Environmental control of the seasonal life cycle of a zoophytophagous mirid, <Emphasis Type="Italic">Adelphocoris triannulatus</Emphasis> (Hemiptera: Miridae)

Adelphocoris triannulatus (Stål) (Heteroptera: Miridae) is a zoophytophagous bug that inhabits grasslands and crop fields, and is widely distributed in eastern Asia. In this study, the seasonal occurrence and environmental factors controlling the seasonal life cycle of this bug were investigated in a local population in southern Kyushu, Japan. When insects were reared under different photoperiods at 20 and 25 °C, they produced diapause or nondiapause eggs depending on the photoperiod: when reared under short-day photoperiods (≤?13 h/day), females laid diapause eggs, whereas they laid nondiapause eggs when reared under long-day photoperiods (≥?14 h/day). Because the developmental duration of the pre-adult stages and maturation (minimum duration of one generation) is estimated to be approximately 30 days at summer temperatures, these results suggest that this bug has a multivoltine seasonal life cycle. In agreement with the laboratory results, only nondiapause eggs were produced until mid-August in the field, and an increasing proportion of diapause eggs were produced from early September. In addition to these findings, field sampling from spring to autumn in 2009 indicated that three or four generations were produced in one year. These results suggest that egg diapause plays a role in preventing nymphs of A. triannulatus from developing in winter, and thereby ensures the survival of the population in adverse seasons.     

Low crowding threshold for induction of sexual reproduction and diapause in a Patagonian rotifer

1. This study investigates the basis and ecological significance of the extremely high propensity for mixis (sexual reproduction) observed in laboratory populations of Brachionus calyciflorus from a temporary pond in Patagonia. 2. Experiments with stem females hatched from resting eggs showed that these females were exclusively amictic but produced mictic daughters even at very low population densities. In six experiments, newly hatched stem females were cultured individually in different volumes (1.5, 12, 40 and 150 mL). The percentage of mictic daughters (mixis ratio) was high in the smaller volumes (e.g. 44–83% in 1.5 mL) and lower in the largest volume (6–21% in 150 mL). A regression analysis combining the data from these experiments showed a highly significant decrease in mixis ratio with volume and indicated that the lowest population density at which mixis still occurs (mixis threshold) would be 3.4 females L^?1 (95% CL 2.9–4.0 females L^?1). This value is considerably lower than mixis thresholds for other rotifers (25–250 females L^?1 for many species and 9000–477 000 females L^?1 for some Spanish Brachionus plicatilis). 3. In three additional experiments, stem females and their amictic daughters were cultured individually in 150 mL. The percentage of mictic daughters produced by these two generations of females was not significantly different, showing that the mixis response to crowding is not inhibited in the stem‐female generation. 4. Laboratory experiments showed that two common predators of the Patagonian B. calyciflorus (the calanoid copepod Parabroteas sarsi and the backswimmer Notonecta vereertbruggheni) each ate 60–70 B. calyciflorus predator^?1 per day and cleared all rotifers from c. 250 mL per day. Thus, a very low mixis threshold and high maximal mixis ratio may ensure production of some resting eggs soon after colonisation of the pond and before complete removal from the plankton. 5. Two laboratory experiments showed that resting eggs of the Patagonian B. calyciflorus hatched at variable rates (28 and 81%) after a brief diapause when kept in the conditions under which they were produced and oviposited (20–21 °C; L: D 16: 8). Early hatching of resting eggs from pond sediment may allow multiple periods of colonisation and resting egg production in a season. This may offset the fitness cost of limited population growth through female parthenogenesis in the face of unpredictable and abrupt risk of extinction because of predators.     

Summer dormancy ensures univoltinism in the predatory bug Picromerus bidens

The seasonal cycle of Picromerus bidens L. (Heteroptera: Pentatomidae) is usually considered to be univoltine with an obligatory winter egg diapause. Seasonal adaptations of the species were studied in the laboratory and in field experiments. When reared under short-day photoperiodic conditions (L12:D12 and L14:D10), all females began to lay eggs synchronously soon after their emergence. However, in the females reared under long-day conditions (L18:D6 and L20:D4) and outdoors in June–July, oviposition was significantly delayed. This delay in reproduction induced by photoperiodic conditions and then spontaneously terminated was considered to be aestivation. Egg batches laid by females in the laboratory and in the field were kept at 25 °C for two months. From 30.8 to 93.8% of batches contained eggs which hatched without cold treatment between day 14 and 60 after oviposition. The proportion of eggs hatched was 17.7 to 20.9% in the short-day regimes, while it was significantly less (5.7 to 6.0%) under long-day conditions. It is concluded that in some eggs diapause is of low intensity and that if under natural conditions the first batches had been laid at the end of June, nymphs would have hatched at least from some eggs during the same season even without cold treatment. Such untimely hatching would have resulted in the death of nymphs and adults unprepared for overwintering. A photoperiodic response which induces aestivation in the early emerging adults in June–August may prevent early oviposition and occurrence of a second generation and thus maintains univoltinism in P. bidens.     

Laboratory rearing method for the stem weevil,Hyperodes bonariensis (Coleoptera: Curculionidae)

The stem weevil, Hyperodes bonariensis Kuschel, was reared from egg to adult for eight generations on an artificial medium. At 26±0.5°C, 55±5% RH, and 12‐h photo‐period, development time from newly hatched larva to adult averaged about 28 days. There were four larval instars lasting about 14 days, followed by prepupal and pupal stages lasting a further 14 days. Oviposition occurred 2 weeks after adult emergence; mated females laid viable eggs. Ovarian diapause in some females was also noted. It is possible to rear about six generations a year under these conditions.     

Factors influencing the induction of diapausing egg production in the calanoid copepod Diaptomus leptopus

Field and laboratory studies were carried out between 1995 and 1997 on four populations of Diaptomus leptopus found in seasonally temporary, occasionally temporary, and a permanent environment to assess the relative importance of photoperiod and temperature regimes versus other proximate local cues in inducing diapause egg production. Patterns of diapausing and subitaneous egg production were determined by observation of individual females bearing clutches that were produced in the field. A laboratory common-garden experiment was performed to assess the effects of four different regimes of temperature and photoperiod on the induction of diapause. Patterns of diapausing egg production differed among ponds: diapause occurred early in the seasonally temporary environment and occurred rarely in the permanent environment. In the common-garden experiment, populations exhibited substantial changes in the onset of diapause when compared to patterns found under field conditions. These results provide indirect evidence that the different populations respond to available cues of environmental change in different ways in nature, or that environmental cues vary among habitats.     

Isolation and characterization ofgrandchildless-like mutants inDrosophila melanogaster

Summary Two temperature-sensitive sex-linkedgrandchildless (gs)-like mutations (gs(1)N26 andgs(1)N441) were induced by ethylmethane sulphonate inDrosophila melanogaster. They complemented each other and mapped at two different loci (1–33.8±0.7 forgs(1)N26 and 1–39.6±1.7 forgs(1)N441), which were not identical to those of any of thegs-like mutants reported in earlier work.Homozygous females of the newly isolated mutants produced eggs that were unable to form pole cells and developed into agametic adults. Competence of the embryos to form pole cells was not restored by wild-type sperm in either mutant; that is, the sterility caused by these mutations is controlled by a maternal effect.Fecundity and fertility ofgs(1)N26 females were low, and their male offspring showed a higher mortality than that of female offspring, causing an abnormal sex ratio. The frequency of agametic progeny was 93.1% and 55.8%, when the female parents were reared at 25° C and 18° C, respectively. In eggs produced by thegs(1)N26 females reared at 25° C, the migration of nuclei to the posterior pole was abnormal, and almost no pole cell formation occurred in these egg. Furthermore, half of these eggs failed to cellularize at the posterior pole. When the females were reared at 18° C, almost all of the eggs underwent complete blastoderm formation, and in half of these blastoderm embryos normal pole cells were formed.In the other mutant,gs(1)N441, the fecundity and fertility of the females were normal. The agametic frequency in the progeny was 70.8% and 18.6% when the female parents were reared at 25° C and 18° C, respectively. In the eggs laid by females reared either at 25° C or at 18° C, the migration of nuclei to the periphery and cellularization proceeded normally; nevertheless, in the majority of the embryos no pole cell formation occured at the stage when nuclei penetrated into the periplasm. When the females were reared at 18° C, some of the embryos from these females formed some round blastoderm cells with cytologically recognizable polar granules and nuclear bodies, which are attributes of pole cells. The temperature sensitive period ofgs(1)N441 was estimated to extend from stage 9 to 13 of King's stages of oogenesis.     

Variable sex ratio strategy of Telenomus heliothidis (Hymenoptera: Scelionidae): adaptation to host and conspecific density

Summary The sex allocation behavior of the solitary egg parasitoid Telenomus heliothidis Ashmead was investigated by examining the response of females reared in isolation and under crowded conditions. Females reared in isolation adjusted their sex ratio with foundress and host number per patch in accordance with the predictions of local mate competition (LMC) theory. However, females did not shift their sex ratio in response to conspecifics foraging on the same host patch or to contact with previously parasitized hosts. Instead, shifts were associated with encounter rate and a sequence of oviposition. Females maintained under crowded conditions responded to host patches much differently. One-day-old females which had lived under crowded conditions for 24 h produced sex ratios similar to those of continuously isolated females. However, females reared under crowded conditions for 7 days consistently produced unbiased sex ratios, and exhibited a different sequence of oviposition. This shift appeared to be due directly to crowding rather than age, oviposition experience or sperm depletion since the effect could be reversed by subsequent isolation.     

Captive rearing of Puget blue butterflies (Icaricia icarioides blackmorei) and implications for conservation

The use of captive rearing to promote recovery of endangered butterflies has substantially increased over the last decade. These programs have the potential to play a significant role in butterfly population recovery, but the effects of captive conditions are poorly understood and rarely are traits of captive individuals assessed relative to traits in their founding populations. To develop rearing protocols and investigate possible effects of captive conditions, we reared Puget blue butterflies (Icaricia icarioides blackmorei), a subspecies closely related to the endangered Fender’s blue (I. i. fenderi) which is limited to Oregon, USA. We reared individuals from two wild populations in Washington, USA to investigate two approaches for egg collection (collect eggs in the wild vs. collect eggs from adult females which were brought to a greenhouse for oviposition) and three diapause environments (in indoor facilities at two independent locations vs. outdoors in enclosures). Survival from egg to adult was similar across all captive groups which survived past diapause and was less than 10%. Captive reared individuals were lighter and had smaller wings and shorter body lengths than their founding populations for both sites. Based on our findings, we recommend that rearing programs compare characteristics of reared individuals to individuals from the founding population to quantify possible effects of captive conditions, diapause individuals in natural environments, and for programs with survival rates similar to rates in the wild, consider alternatives to augment declining populations and reintroduce historic ones.     

Establishment of in vitro plants, cell and tissue cultures from Oldenlandia affinis for the production of cyclic peptides

In vitro cultured plants from Oldenlandia affinis were established from seeds and grown on a hormone-free medium. In vitro plants produced the cyclic peptide kalata B1 in concentrations of 0.67 mg g⁻¹ dry weight after growth of 30 days. This was approximately 50% of the concentration analysed in green house plants (shoot tips), where different concentrations have been determined in leaves (1.82 mg g⁻¹), shoot tips (1.36 mg g⁻¹), stems (0.36 mg g⁻¹), and in flowers (0.16 mg g⁻¹). Callus and cell suspension cultures could be initiated from aseptic root, stem and leaf explants of O. affinis seedlings and plants. Different light intensities were shown to affect culture growth as well as chlorophyll synthesis. The friable callus was then used for the establishment of a cell suspension culture. Fresh and dry weight measurements showed that growth was optimal on MS medium supplemented with 0.4 mg l⁻¹ 2,4-dichlorophenoxyacetic acid (2,4-d). Leaf suspensions cultured on this medium showed a 4-fold increase of biomass by the first week of incubation. No quantifiable amounts of kalata B1 were produced under these conditions. Morphological differentiation seems to be essential for cyclic peptide production. Therefore, several undifferentiated as well as organised cell lines of O. affinis have been developed. These cell lines will constitute a worthwhile starting point for the optimisation of kalata B1 synthesis in liquid media to the objective of producing cyclic peptides under controlled and defined conditions in bioreactors.     

Egg load of lab-cultured <Emphasis Type="Italic">Anaphes iole</Emphasis> and effects of mate presence and exposure time on load depletion

Anaphes iole Girault (Hymenoptera: Mymaridae) is a solitary egg parasitoid of Lygus bugs (Heteroptera: Miridae) in North America. This research considered factors that might impact the egg load of lab-cultured A .iole females, reared from Lygus hesperus Knight egg patches. The following hypotheses were tested: (1) egg load was related to body size and not affected by female age, and (2) egg load depletion was not affected by mate presence and time (in days) that females were exposed to host patches. Initial egg load averaged 48 mature eggs and no immature eggs were detected in the ovarioles of dissected females. Egg load was neither related to body size (hind tibia or forewing length) nor affected significantly by age (0, 1 or 2days old honey-fed females). Mate presence (females with or without males) and exposure time (1, 3 or 5days on the same host patch) had no effect on egg load depletion. Females usually depleted most of their egg load within 24h. From 86 to 92% of females contained less than six mature eggs and no immature eggs after 1, 3 or 5days of exposure to host patches. The results of this study suggest that A. iole females are certainly pro-ovigenic and initial egg load does not correlate with body size or age. Since mated and unmated females deplete most of their egg load in 24h, time-efficient production of progeny may result when ovipositing parasitoids are exposed to suitable hosts for just a few days.The United States Government has the right to retain a non-exclusive, royalty-free license in and to any copyright of this article. This article reports the results of research only. Mention of a commercial or proprietary product does not constitute an endorsement of the product by the United States Department of Agriculture.