Foraging strategy of wasps - optimisation of intake rate or energetic efficiency?

In endothermic wasps, foraging is an expensive activity. To maximise the benefit for the colony, wasps can optimise either the intake rate or energetic efficiency of a foraging trip. We investigated the foraging behaviour of vespine wasps under variable environmental and reward conditions. We trained them to forage for 0.5 mol l-1 sucrose solution from an artificial flower in a flow-through respiratory measurement chamber, and simultaneously measured their body temperature using infrared thermography to investigate interactions between thermoregulation and energetics. Measurement of carbon dioxide release (for energetic calculations) and load mass enabled the direct determination of foraging efficiency. An unlimited reward increased the wasps' energetic effort to increase the suction speed through high body temperatures. With reduced reward (limited flow), when an increased body temperature could not increase suction speed, the wasps decreased their body temperature to reduce the metabolic effort. Solar heat gain was used differently, either to increase body temperature without additional metabolic effort or to save energy. The foraging efficiency was mainly determined by the flow rate, ambient temperature and solar heat gain. In shade, an unlimited sucrose flow and a high ambient temperature yielded the highest energetic benefit. A limited flow reduced foraging efficiency in the shade, but only partly in sunshine. Solar radiation boosted the efficiency at all reward rates. Wasps responded flexibly to varying reward conditions by maximising intake rate at unlimited flow and switching to the optimisation of foraging efficiency when the intake rate could not be enhanced due to a limited flow rate.

What do foraging wasps optimize in a variable environment, energy investment or body temperature?

Vespine wasps (Vespula sp.) are endowed with a pronounced ability of endothermic heat production. To show how they balance energetics and thermoregulation under variable environmental conditions, we measured the body temperature and respiration of sucrose foragers (1.5 M, unlimited flow) under variable ambient temperature (T a = 20-35 °C) and solar radiation (20-570 W m(-2)). Results revealed a graduated balancing of metabolic efforts with thermoregulatory needs. The thoracic temperature in the shade depended on ambient temperature, increasing from ~37 to 39 °C. However, wasps used solar heat gain to regulate their thorax temperature at a rather high level at low T a (mean T thorax ~ 39 °C). Only at high T a they used solar heat to reduce their metabolic rate remarkably. A high body temperature accelerated the suction speed and shortened foraging time. As the costs of foraging strongly depended on duration, the efficiency could be significantly increased with a high body temperature. Heat gain from solar radiation enabled the wasps to enhance foraging efficiency at high ambient temperature (T a = 30 °C) by up to 63 %. The well-balanced change of economic strategies in response to environmental conditions minimized costs of foraging and optimized energetic efficiency.

Performance of Psyttalia humilis (Hymenoptera: Braconidae) reared from irradiated host on olive fruit fly (Diptera: Tephritidae) in California.

The parasitoid Psyttalia humilis (Silvestri) was reared on Mediterranean fruit fly, Ceratitis capitata (Wiedemann), third instars irradiated at 0-70 Gy at the USDA, APHIS, PPQ, Moscamed biological control laboratory in San Miguel Petapa, Guatemala, and shipped to the USDA, ARS, Parlier, CA. Irradiation dose did not affect the parasitoid's offspring sex ratio (53-62% females), percentage of unemerged adults (12-34%), number of progeny produced per female (1.4-1.8), and parasitism (19-24%). Host irradiation dose had no significant effect on the forewing length of female P. humilis and its parasitism on olive fruit fly, Bactrocera oleae (Rossi) and offspring sex ratio, but dissection of 1-wk-old female parasitoids reared from hosts irradiated with 70 Gy had a significantly lower number of mature eggs than females from nonirradiated hosts. Longevity of P. humilis adults decreased with increased temperature from 15 to 35°C, regardless of food provisions, gender, and host irradiation dose. Females survived 37-49 d at 15°C with water and food, and only 1-2 d at 35°C without food, whereas males lived shorter than females at all temperatures and food combinations tested. Adult P. humilis reared from fertile C. capitata and aspirated for dispensing in cups lived significantly longer after shipment than those specimens chilled and dispensed by weight. At 21 and 32°C, 50% of parasitoids departed release cages after 180 and 30 min, respectively, but none departed at 12°C. Thirteen shipments of P. humilis (2,980-21,922 parasitoids per shipment) were received between September and December 2009, and seven shipments (7,502-22,560 parasitoids per shipment) were received between October and December 2010 from San Miguel Petapa, Guatemala. Daily number of olive fruit fly adult and percentage female trap captures ranged <1-19 and 8-58% in 2009, and <1-11 and 0-42% in 2010, respectively. The number of parasitoids released ranged 848-12,257 in 2009 and 3,675-11,154 in 2010. Percentage parasitism of olive fruit fly third instars at all locations ranged 0-9% in 2009 and 0-36% in 2010.

Circadian rhythms of adult emergence and activity but not eclosion in males of the parasitic wasp Nasonia vitripennis.

An endogenous circadian system is responsible for the rhythms observed in many physiological and behavioural traits in most organisms. In insects, the circadian system controls the periodicity of eclosion, egg-laying, locomotor and mating activity. The parasitoid wasp Nasonia vitripennis has been extensively used to study the role of the circadian system in photoperiodism. In this study, behavioural activities expected to be under the control of the endogenous circadian system were characterized in Nasonia. Male emergence from the host puparium is rhythmic under light-darkness conditions while eclosion from the own pupal integument is not rhythmic but continuous. Following entrainment in light-dark conditions, males show robust free-running circadian activity rhythms with a period (tau, tau) of approximately 25.6h in constant darkness. While the endogenous circadian system is enough to trigger male emergence in Nasonia, light seems to have a modulatory effect: when present it induces more males to emerge. Our results add to the understanding of chronobiological phenotypes in insects and provide a basis towards the molecular characterization of the endogenous circadian system in Nasonia.

Some liver functions in the Oriental hornet (Vespa orientalis) are performed in its cuticle: exposure to UV light influences these activities.

The Oriental hornet Vespa orientalis (Hymenoptera, Vespinae) coordinates its daily activities (e.g. flights out of the nest associated with digging activities and removal of the dug soil from the nest) with the amount of insolation. Thus, the stronger the insolation, the more intense the flight activity and vise versa. The hornet's cuticle bears a few yellow stripes interposed among brown parts of the gastral cuticle. These yellow stripes are composed of two elements, namely, a transparent cuticle and underneath it a layer of yellow granules. When the hornets are exposed to UV light, the layer containing the yellow granules is less active than in hornets kept in the dark. This diminished activity entails a lower production of glucose as well as of several enzymes prevalent also in the liver of mammals, like creatine kinase, alanine aminotransferase, aspartate transaminase. Thus solar irradiation stimulates and produces a change in the metabolic activities of the hornet. The fact that hornets link their flight activity with the insolation leads us to speculate that the sun contributes energetically to the hornet's activity.

Impact of environmental stress on aphid clonal resistance to parasitoids: Role of Hamiltonella defensa bacterial symbiosis in association with a new facultative symbiont of the pea aphid.

Resistance to endoparasitoids in aphids involves complex interactions between insect and microbial players. It is now generally accepted that the facultative bacterial symbiont Hamiltonella defensa of the pea aphid Acyrthosiphon pisum is implicated in its resistance to the parasitoid Aphidius ervi. It has also been shown that heat negatively affects pea aphid resistance, suggesting the thermosensitivity of its defensive symbiosis. Here we examined the effects of heat and UV-B on the resistance of A. pisum to A. ervi and we relate its stability under heat stress to different facultative bacterial symbionts hosted by the aphid. For six A. pisum clones harboring four different facultative symbiont associations, the impact of heat and UV-B was measured on their ability to resist A. ervi parasitism under controlled conditions. The results revealed that temperature strongly affected resistance, while UV-B did not. As previously shown, highly resistant A. pisum clones singly infected with H. defensa became more susceptible to parasitism after exposure to heat. Interestingly, clones that were superinfected with H. defensa in association with a newly discovered facultative symbiont, referred to as PAXS (pea aphid X-type symbiont), not only remained highly resistant under heat stress, but also expressed previously unknown, very precocious resistance to A. ervi compared to clones with H. defensa alone. The prevalence of dual symbiosis involving PAXS and H. defensa in local aphid populations suggests its importance in protecting aphid immunity to parasitoids under abiotic stress.

Hornet flight activity and its correlation with UVB radiation, temperature and relative humidity.

During the active season, extending from June to October, hornets emerge from their nest in the field in all the daytime hours. In the beginning of the season, when the number of workers is relatively small, the number of exits from the nest is fairly uniform numerically throughout the day. However, with the increase in hornet population from July onwards, the number of workers emerging from the nest entrance around noon (1100-1300 h) is by 1-2 orders of magnitude greater than the number of those emerging in the morning or evening hours. This disparity persists till September or October, at which time the workers revert to behave as in the beginning of the season. It appears, therefore, that in this period hornet activities outside the nest are coordinated with the meteorological conditions, and in this regard, the highest correlation is with the ultra violet B (UVB) radiation level and to a lesser extent with the temperature. Presumably, also, the greater noon-hour activity in the nests of hornets in the field stems from the digging hornets benefiting from the greater availability of solar energy at noon, mainly that of UVB radiation. We assume that the hornets are able to utilize the UVB radiation, but what part of their body is "absorbing" the UVB energy is still a matter of further investigation.

[Destabilization of the cardiac function of an insect by a low-frequency electric field].

The effect of high-intensity low-frequency electric field on the functioning of the heart of an insect was estimated from electrocardiogram. It was found that electric field causes a disturbance of the cardiac function. Its stressing activity is mainly related to the excitation of the insect by induced currents whose exciting action is enhanced by vibration in trichoid sensillas and antennas.

Hornet flight is generated by solar energy: UV irradiation counteracts anaesthetic effects.

The Oriental hornet Vespa orientalis (Hymenoptera, Vespinae) flies outside its nest only in the daytime and never in the dark. Oriental hornets can be anaesthetized by means of diethyl ether, following which they awake spontaneously within about > or =1 h. However, when the anaesthetized hornets are exposed to direct irradiation they awake much faster and immediately fly out and this is within the temperature range for their normal activities (20-40 degrees C). Light exerts an effect not only on intact hornets, but also on their main body parts, be they the head, thorax or gaster, or a torso without a head or without a gaster. These body parts also 'awake' from ether anaesthesia when illuminated, especially by ultraviolet (UV) B or shorter light wavelengths, but a body without a gaster awakes much later than normal. As for flight activity, headless hornets do not fly at all, while hornets devoid of a gaster flutter their wings when illuminated but are incapable of rising in the air. When vespan cuticular regions are coated with paint that blocks the passage of light, such as UVB blockers, or with white Tipp-Ex, the awakening from anaesthesia is delayed. The vespan cuticle evinces photovoltaic activity, even in dead specimens, so that upon exposure to light it is possible to record levels of 30-180 mV, both in a wakeful hornet and in one that has undergone anaesthesia. We conjecture that in the awake hornet the voltage that its cuticle generates under the influence of light is transported within the cuticular interlamellar membranes to be utilized as flight energy by the muscles attached to them; however, in the narcotized hornet, the diethyl ether apparently decreases or disrupts membranal order, blocking any ionic channel activity and thereby inducing the anaesthesia. Presumably, this state of phase transition is remedied spontaneously, albeit in a lengthy process, or else the light energy in UV speeds up restoration of the ionic channel activity and the synaptic transmission and thereby contributes to a more rapid awakening. In this regard, the hornet gaster is an extensive region for absorbing light, which explains why its absence considerably delays the awakening from anaesthesia.

Photovoltaic effects in hornets are correlated with the time of day.

This study deals with voltage values recorded off the cuticle of live specimens of the Oriental hornet Vespa orientalis (Hymenoptera, Vespidae). The relevant measurements were taken between the two tips of their bodies at various hours of the day and were made on a total of 90 worker hornets. Recorded voltage values varied within a range of 60-180 mV, with the lower values measured during the morning hours and the afternoon and the highest values during the noon hours. Measurements were made by direct contact of the electrodes with the hornet cuticle and did not prove lethal to the measured specimens. An additional 60 live hornets were measured in the same fashion but in the dark. The values recorded in the dark varied between 40 and 70 mV and displayed considerable fluctuations but were not found to be dependent on the time of measurement. The distribution of the voltage values in hornets measured at various hours in the daytime closely resembled that of the global radiations (in W/m2) on the same days the measurements were taken.

Photo-induced arousal response by hornets.

When an Oriental hornet Vespa orientalis is subjected to ether anesthesia and then exposed to ultraviolet A light (UVAL) (at a wavelength of 366 nm), it commences showing signs of awakening by starting to move its limbs. While in the process of waking the voltage on its body surface surges sharply from 17-180 mV (median = 71.0) to a level of 93-570 mV (median = 327.5). This elevated level is maintained for several minutes but subsequently drops sharply to starting level. The increase in voltage is throughout accompanied by fluttering of the wings and movements of the legs, as well as attempts to extricate itself from the bindings to the electrodes. These movements by the awakening hornet persist for several minutes even after the irradiation source is turned off but shortly after the switch-off the hornet lapses into sleep again. The described scenario is generally similar in worker, queen and drone hornets, and may even occur in decapitated specimens. The same type of awakening can be repeated in the same fashion after a while, but then the increase in voltage will be smaller than the first time. Continuous UV irradiation of an anesthetized hornet results in a generalized and protracted awakening which, however, is significantly shorter than in a hornet left anesthetized in the dark.

Hornet cuticle: effects of short-term UV irradiation.

Effects of short-term UV irradiation were investigated on various cuticular parts of workers and queens of the Oriental hornet, to wit: brown strip, yellow strip and wing. On each preparation of the afore-mentioned, a reading of the relative optical density (ROD) was taken prior to, immediately following, and 15-30 minutes after its irradiation as compared to white light irradiation. The results showed that brief UV irradiation causes changes in the ROD of hornet cuticle, and that these changes in ROD are different in brown than in yellow cuticle. Those in yellow strip are induced by the presence or absence of the active yellow pigment, whose quantity in worker cuticle is different than in queen cuticle, probably due to the various activities in which they are involved during the active season.

Hornet peak flight activity is correlated with solar UV radiation: a multi-annual survey.

This study deals with the effect which solar irradiation of short wavelength, particularly ultraviolet (UV), exerts on the activities of hornets. The findings are based on multi-annual observations carried out during the years 1985, 1989 and 1998 on hornet nests in the field. At the peak of UV radiation, which occurs at noon, hornet activity is greater by 1-2 orders of magnitude than that during the morning or evening hours. The main visible hornet activity appears to be the removal of soil particles from the nest so as to enlarge its volume, enable the building of additional combs and also increase the size of existing combs. Hornet flight during peak insolation hours is characterized by its briefness (5-20 seconds only) and brevity (to distances of 5-10 meters only) as compared to flights at other hours of the day. These prolonged, multi-annual observations lead to the conclusion that hornets are capable of converting the energy of UV radiation into a form amenable to metabolic usage. In this respect the hornet cuticle behaves as a thermophotovoltaic device, i.e., a semiconductor diode that converts photons radiating from the sunlight into electrical energy.

Thermophotovoltaic (TPV) properties of hornet cuticle as dependent on relative humidity.

This paper deals with the thermophotovoltaic (TPV) properties of the cuticle of the Oriental hornet as assessed over time under different regimens of relative humidity (RH). The tests were run at two levels of RH, namely, 30% vs. 90%. Each experiment entailed measuring the cuticular voltage and current in the dark as compared to under illumination (white light = 700 Lux), and at a temperature range of 20-30 degrees C. It was found that increase in the RH level boosts the current values by 2-3 orders of magnitude; contrariwise, the voltage values rise by about three times with drop in the RH. At high RH, the changes in current become rhythmical and each cycle of warming-cooling assumes a distinctly cyclic pattern. Under illumination, the current decreases, the polarity reverses and the resistance increases. The obtained results are describable by a model of electric conductance upon a surface, in this case the hornet cuticle; the findings are also discussed and compared with similar phenomena recorded from other substances possessing the properties of organic semiconductors.

Dispersal and parasitizing abilities of Eupelmus vuilleti (Hymenoptera: Eupelmidae) within a column of cowpea seeds.

Eupelmus vuilleti (Crawford) is an ectoparasitoid of the seed-eating beetle Bruchidius atrolineatus (Pic), which is an important pest of stored cowpea, Vigna unguiculata Walp, seeds in West Africa. Herein, we investigated the dispersal abilities of females within columns of seeds to assess the potential of E. vuilleti as a biological control agent of bruchids in cowpea granaries. The influence of host presence together with the 2 abiotic factors light and gravity on parasitoid movement and parasitization efficiency were analyzed. E. vuilleti females were able to travel through large seed masses and parasitize hosts located at the end of the seed column opposed to their introduction zone. Parasitoid movement was stimulated by light. E. vuilleti females exhibited a negative geotropism. Females introduced at the bottom of the seed column dispersed more and parasitized more hosts than females introduced at the top. Host presence had some influence on the dispersal of the parasitoids within the seed column at a host density of 10 infested seeds for 16,000-18,000 uninfested seeds. This depended on female introduction zone because gravity was the major factor influencing dispersal. The possible applications of these results for biological control of bruchids in cowpea granaries are discussed.

Duration of hornet sleep induced by ether anesthesia is curtailed by exposure to sun or UV irradiation.

Irradiation of hornets anesthetized by diethyl ether curtails their sleep duration from the ordinary 20-30 min to a mere 2-3 min. This effect on sleep is dependent on the intensity of the sun irradiation or, when exposed to monochromatic UV at 366 nm, on the duration of the irradiation. Of the various hornet cuticular areas of the body, the yellow cuticular areas of the abdominal segments are the most sensitive to the treatment. We assume that the cuticles of both live and dead hornets act as a converter of light to electric energy.

Deletion analysis of the selfish B chromosome, Paternal Sex Ratio (PSR), in the parasitic wasp Nasonia vitripennis.

Paternal Sex Ratio (PSR) is a "selfish" B chromosome in the parasitoid wasp Nasonia vitripennis. It is transmitted via sperm, but causes supercondensation and destruction of the paternal chromosomes in early fertilized eggs. Because this wasp has haplodiploid sex determination, the effect of PSR is to convert diploid (female) eggs into haploid (male) eggs that carry PSR. Characterizing its genetic structure is a first step toward understanding mechanisms of PSR action. The chromosome is largely heterochromatic and contains several tandemly repeated DNA sequences that are not present on the autosomes. A deletion analysis of PSR was performed to investigate organization of repeats and location of functional domains causing paternal chromosome destruction. Deletion profiles using probes to PSR-specific repetitive DNA indicate that most repeats are organized in blocks on the chromosome. This study shows that the functional domains of PSR can be deleted, resulting in nonfunctional PSR chromosomes that are transmitted to daughters. A functional domain may be linked with the psr22 repeat, but function may also depend on abundance of PSR-specific repeats on the chromosome. It is hypothesized that the repeats act as a "sink" for a product required for proper paternal chromosome processing. Almost all deletion chromosomes remained either functional of nonfunctional in subsequent generations following their creation. One chromosome was exceptional in that it reverted from nonfunctionality to functionality in one lineage. Transmission rates of nonfunctional deletion chromosomes were high through haploid males, but low through diploid females.

The influence of X-rays on reproduction in Pseudeucoila bochei (Hymenoptera Cynipidae), a parasite of Drosophila melanogaster.

The parasitic wasp Pseudeucoila bochei was the subject in experiments designed to detect a reliable method of mutation induction. Radiation doses of 360, 720, 1080, 1440, 1800 and 2160 R were given to pupae 15 or 18 days old or to freshly eclosed male and female wasps. The irradiation did not influence the survival of pupae. Dose-action responses were established and appeared to be dependent on the stage of development of the wasps at the moment of irradiation. The production of sons by virgin females showed that the germ cells of the adults were the most sensitive to irradiation. In contrast, the sperms of the male wasps irradiated as adults were the most resistant. Irradiation of males as pupae with 1880 or 2160 R on the 15th day or as adults led to a high frequency of amle progeny. The elimination of sperm cells from fertilization is discussed. Comparisons with Habrobracon, Nasonia and Drosophila showed that the germ cells of Pseudeucoila are more sensitive to irradiation. In the progeny from irradiated females and males 2 types of mutant were found, with yellow eye colour and with aberrations in the antennae.

Biological responses of Habrobracon to spaceflight.

Since the interaction of the parasitic wasp Habrobracon with the space environment could not be prejudged, we decided to test approximately 30 different parameters of a genetic, mutational, biochemical, behavioral, and physiological character in the one spaceflight we had at our disposal. These parameters were examined at six different exposures of gamma-radiation (including 0 dose) in flight, resulting in about 180 different endpoints in all. The most profound effects of spaceflight in conjunction with radiation were decreased hatchability and enhanced fecundity of eggs exposed to spaceflight at different stages of oogenesis. The interpretation we favor is that these two endpoints are reflections of chromosomal non-disjunction in the former case and inhibition of cell division in the latter. Our most comprehensive study of mutagenesis was on sperm, where dominant lethality, recessive lethality, translocations, and visible mutations were assayed; the only effect found was a threefold enhancement of the recessive lethal mutation frequency in the non-irradiated sperm in the orbited Habrobracon males. Behavioral and biochemical differences were found. Mating activity of orbited males was severely disrupted and xanthine dehydrogenase activity was sharply decreased in the irradiated flight animals, an unexpected observation. Postflight experiments were like the ground-based control experiments in all aspects but one. Under conditions of vibration similar to those encountered during the launch and re-entry, the mutation frequency in the sperm increased by a factor of three over that of the non-vibrated control.

Biosatellite II--physiological and somatic effects on insects.

Interesting physiological and somatic effects of space flight have been observed in a variety of insects on board Biosatellite II. Wasps, beetles and fruit flies have demonstrated positive effects of the space flight alone or in combination with irradiation. In the flour beetle, Tribolium confusum, the incidence of a developmental wing abnormality was measured. This abnormality, which mimics the mutation known as "split", was significantly increased in beetles which were flown as young pupae, pre-irradiated with X rays to bring them into their sensitive dose range and gamma irradiated during flight. Wing abnormalities increased from a ground value of 29.9 per cent to a value of 44.8 per cent. The length of the pupal period was also measured. Although there was a significant increase of the flight pupal period over that of the ground control, the most likely explanation seems to be the temperature drop of the flight samples between separation and retrieval of the flight capsule. In the parasitic wasp, Habrobracon juglandis, flown in the adult stage, several interesting results have been obtained in terms of reproductive performance, life span and enzyme activity. Reproductive performance, measured by the average number of eggs laid/female/day, was unaffected by flight for control unirradiated wasps. However, for females X rayed with 2000 R prior to flight, the characteristic depression of egg-laying at 10 days post-irradiation was negligible in the flight animals. A more pronounced effect was observed in those females which received the chronic 2667 R gamma ray dose in flight; egg production actually progressively increased to twice the level of the ground sample. It was observed that males were disoriented in their mating behavior for 2 days after the flight. When the dose level data was pooled, the life span of females was found to be significantly longer in the flight group. Upon analysis of the activity of the enzyme, xanthine dehydrogenase (XDH), adult flight males were found to have a significantly decreased XDH activity compared with ground controls. No significant difference was found between female F1 progeny of flight and ground males. XDH activity was also analyzed in F1 progeny of Drosophila melanogaster, orbited as adults and larvae. For the F1 male progeny of adult males, there was a significantly depressed average XDH activity compared with ground controls for the highest irradiation dose. For F1 male progeny of flight larvae, there was a statistically decrease of XDH activity only between the off-spring of irradiated flight and ground larvae. Further somatic changes were observed in Drosophila larvae, in which chromosome aberrations were studied.