Resistance of apple trees to Cydia pomonella egg-laying due to leaf surface metabolites

During host plant selection and particularly after alighting on a plant, chemical cues from the plant surface influence an insect's acceptance of the plant and, subsequently, its egg‐laying behaviour. Primary metabolites in the phylloplane may be more important than hitherto known. We have shown that soluble carbohydrates, such as glucose, fructose, and sucrose, and sugar alcohols, such as sorbitol, quebrachitol, and myo‐inositol, can be detected by insects after contacting the plant and that they positively influence egg‐laying of the codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae), on apple trees. We addressed the question whether a lack of these substances could also explain apple tree resistance to C. pomonella in terms of reduced egg‐laying. Leaf surface washings were collected in an apple orchard by spraying water on the resistant cultivar X65‐11 and on the susceptible cultivar P5R50A4. The washings were tested on a nylon cloth on isolated females under no‐choice conditions. The washings were analysed and synthetic blends, each consisting of the six metabolites in the proportions established in the leaf surface washings of both cultivars, were then tested for their effect on egg‐laying of C. pomonella. Dose–response egg‐laying tests were carried out on substrates impregnated with the X65‐11 leaf surface blend at 1, 100, 1 000, and 10 000 times the natural dose. Egg‐laying behaviour in the bioassays with leaf surface washings of both cultivars closely resembled egg‐laying in the orchard. Washings of P5R50A4 stimulated egg‐laying to a greater extent than those of X65‐11 and the water control. Synthetic blends reduced substrate acceptance and egg‐laying, compared to the washings of X65‐11. Ratios between components within the blend are responsible for this resistance. In conclusion, quantities and ratios of the six primary metabolites found on the leaf surface may influence host preference of C. pomonella as well as their egg‐laying behaviour, thus they may play a role in the trees’ resistance to the codling moth.     

Egg-laying experience and acceptance of parasitized hosts by the parasitoid,Leptopilina heterotoma (Hymenoptera: Eucoilidae)

The influence of egg-laying experience on the response of females of the eucoilid parasitoid,Leptopilina heterotoma, to parasitized and unparasitizedDrosophila melanogaster host larvae was examined under more controlled conditions than those used in past studies. In laboratory assays, we precisely manipulated both the number of eggs laid by females and the kind of larvae (parasitized versus unparasitized) in which the eggs were laid. We found that the tendency to avoid laying eggs in parasitized hosts depended markedly on whether or not eggs had been laid previously, but depended little on whether those eggs had been laid in parasitized or unparasitized hosts. The observed effect of general egg-laying experience on avoidance of parasitized hosts may reflect responses to either changes in the wasp's internal state (perhaps, changes in egg load) or changes in the wasp's neural representation of the external environment (such as those presumed to occur during learning). In light of these results, we offer a tentative reinterpretation of several earlier studies.     

Temperature and egg-laying experience influence breeding performance of captive female houbara bustards

The influence of climate, age and egg-laying experience on reproductive performance (propensity, fecundity, timing) of a flock of captive houbara bustards was determined. The flock, established in Saudi Arabia for reintroduction purposes, was housed in outdoor pens, and fed food and water ad libitum. Birds were artificially inseminated when they came into breeding condition and eggs were removed soon after laying. Winter temperatures, rather than rainfall, synchronized reproduction in the captive flock. Cooler winter temperatures led to higher proportions of females laying, an earlier start to laying, and greater numbers of eggs laid per season. The first significant rainfall of the season stimulated some females to lay their last egg. Age per se had little effect on breeding performance independent of egg-laying experience. Females with no previous egg-laying experience laid fewer eggs than similarly aged females with previous experience. This difference, in combination with a pattern of increasing recruitment into the breeding population between 1 and 5 years of age, led to improved breeding performance in the population with age. The inferior performance of first-time egg-layers, despite ad libitum food, indicates restraint in the onset of breeding, which may be an adaptation for optimizing life-time fitness.     

Manipulation of laying effort reveals habitat-specific variation in egg production constraints in Great Tits (Parus major)

It has been suggested that a bird's clutch size is not limited by the amount of resources available at the time of laying but that differences in the availability of food for nestlings is the ultimate underlying factor determining spatio-temporal variations in clutch size. However, habitat-related variations in egg production ability has yet to be investigated explicitly. We studied the breeding of Great Tits Parus major in deciduous and coniferous forests in the same area. The sizes of both the clutches and the eggs were, on average, larger in the former habitat than in the latter. A number of females were induced to lay more eggs than usual by removing four eggs from designated experimental clutches early in the laying period. These manipulated females laid approximately one egg more than control females, with the number of additional eggs laid not differing between the habitats. However, in both study years the relative size of the extra eggs – relative to the mean size of earlier laid eggs of the same clutch – was smaller in the coniferous habitat than in the deciduous habitat, while there was no habitat-related difference in the relative size of the last-laid eggs of control clutches. This result indicates that some form of proximate limitation during egg-laying period can contribute to the relatively small clutches and eggs in the coniferous habitat. Our results emphasize the need to take egg production costs into account when attempting to account for spatial variation in the reproductive behaviour of birds.     

广聚萤叶甲成虫产卵行为及其产卵部位的选择性

【目的】明确广聚萤叶甲成虫产卵行为及其产卵部位的选择性。【方法】在室内条件下,对广聚萤叶甲成虫交配及产卵的系列行为、产卵场所选择、不同部位豚草植株叶片叶绿素b的含量进行了观察和测定:(1)将1对成虫放到养虫笼内的一株豚草上,观察交配时间,记录产卵数量、前后2粒卵之间的产卵时间间隔;(2)在均匀分为5部分(0～10、11～20、21～30、31～40和41～50 cm)的豚草植株上,随机放置10对成虫,观察雌虫对于产卵场所的选择。(3)将上述5个部位的豚草叶片通过丙酮匀浆法处理,用紫外分光光度计测定其在645和663 nm的吸光值,计算叶绿素b含量。【结果】广聚萤叶甲成虫完成一次成功交配平均需96.09 min。雌虫一般需45 min的时间来寻找其适应的产卵场所,在产卵过程中,成虫习惯将卵产于叶片背面,雌虫喜欢用口器来清理刚产下的卵粒。在一株50 cm高的豚草植株上,雌虫喜欢将卵产在植株中部21～30 cm和中上部31～40cm的叶片上(从下往上划分)。卵块数量和豚草不同部位叶片叶绿素b含量呈显著的正相关性。【结论】广聚萤叶甲成虫喜欢产卵在叶绿素b含量较高的叶片背面,可能以视觉识别叶片颜色来选择和定位产卵场所。     

Geographic differences on accumulation of sugars and polyols in locust eggs in response to cold acclimation

The accumulation of low molecular weight sugars and polyols is one of major mechanisms hypothesized to increase cold tolerance in overwintering insects. But little is known about whether these sugars and polyols are involved in geographic variation of cold tolerance. In this study, we investigated accumulation patterns of eight low molecular weight sugars and polyols of eggs in tropical and temperate populations of the migratory locust, which exhibits between-population variation in cold tolerance, in response to cold acclimation (5, 0 and −5 °C). Excluding erythritol, the other seven carbohydrates were identified as possible cryoprotectants in locust eggs. Basal maximal and minimal concentrations were 45 μg/g wet weight for trehalose and 0.59 μg/g wet weight for glycerol. Most sugars and polyols were elevated after a −5 °C exposure. In a tropical population, fructose, glucose, sorbitol and myo-inositol were significantly accumulated by low temperature treatments, but glycerol was not. In the temperate population, glycerol, glucose, mannitol, sorbitol, myo-inositol were significantly accumulated but trehalose did not increase. Our results suggest different accumulation patterns of these carbohydrates of locust eggs between tropical and temperate populations and highlighted possible roles for them in geographic variation of cold tolerance in the migratory locust.     

薜荔榕小蜂产卵和传粉行为

薜荔榕小蜂Blastophaga pumilae是薜荔的传粉昆虫,栖息于薜荔隐头花序中,在其生物学行为中以钻入花序产卵或传粉最为关键。研究表明：薜荔榕小蜂钻入花序平均费时69.48 min,标准差13.26 min;伤残程度很高并能在伤残的情况下完成产卵或传粉;卵的尾丝能起标识作用避免重复产卵;进入雌花序的小蜂具逃离花序的行为但不能实现;1只小蜂平均产528个卵或为1 072.4朵雌花授粉。     

Interaction of a population of the black-kneed capsid, Blepharidopterus angulatus, and its prey, the lime aphid

Abstract.

• 1 The population dynamics of Blepharidopterus angulatus (Heteroptera: Miridae), preying on the lime aphid, were investigated from 1965 to 1974. Numbers increased or remained stable until 1969, then declined, at least intermittently. K-factor analysis showed that the key factor was k₅₊₆(loss of adult females, eggs and newly-hatched nymphs).
• 2 Detailed study of numbers and mortalities from 1968 to 1971 showed that loss of adult females was the largest part of k₅₊₆. Studies of flight activity showed most females probably emigrated from the trees before laying eggs, but k₅₊₆ was not related to aphid numbers at the time of peak migration. However, variation in k₅₊₆ depended on the length of time that aphid numbers remained favourable for egg-laying.
• 3 Because of the consistently high emigration of adult females, B.anguhtus numbers increased on average by only 10% when aphid numbers favoured egg-laying. Nett emigration was usually high, probably because the trees were isolated or in rows, with few sources of immigrants to the South-West (direction of prevailing wind). Since the population declined by 85% in years when aphids favoured egg-laying for ≤ 20 days, recovery would be slow after a bad year. Since 1969, 1971 and 1973 were all bad years, this explains the intermittent decline in B.angulatus numbers after 1969.
• 4 Simulation showed that neither increased numbers nor faster population growth of B.angulatus would enhance stability of predator and prey populations, but would lead to over-exploitation of the latter. Given its low observed rate of increase, it is unlikely that B.angulatus could persist on lime at levels much above those recorded.

     

Errors in egg-laying by female Common Cuckoo Cuculus canorus in nests of its common host

Dozens of studies have documented that brood parasites are well adapted to a brood parasitic lifestyle but not all parasitism events are successful. Co-evolution between brood parasites and their hosts is a dynamic process so it is reasonable to expect that a female brood parasite may commit errors during egg deposition by laying her eggs outside the laying period of the host, with consequent impacts on her fitness. Using an extensive dataset from a long-term study, we evaluated egg-laying patterns and errors related to the timing of egg-laying in the Common Cuckoo Cuculus canorus (hereafter ‘Cuckoo’). Specifically, we tested whether the Cuckoo avoids laying before or on the day of host clutch initiation to reduce the risk of rejection of parasitic eggs, whether laying errors will be more frequent in periods with a lack of active host nests, and whether the laying errors will be more frequent in periods with intense Cuckoo parasitism and a consequent lack of suitable host nests. We found that about one-third of Cuckoo eggs were laid on the host clutch initiation day or 1 day before, and the percentage of Cuckoo eggs laid decreased thereafter. Surprisingly, the probability of Cuckoo egg acceptance by the hosts was not affected by the egg-laying stage of the host clutch. Errors in the timing of egg-laying with fatal consequences (i.e. those precluding Cuckoo hatching because of laying in incubated or deserted clutches) were recorded in about 5% of cases. Only laying date of a Cuckoo egg had a significant effect on the probability of errors, which increased during the breeding season. This may be related to the higher number of deserted and incubated host nests at the site at the end of the breeding season. Errors in egg-laying may be attributed to young and inexperienced females but also impaired body condition or intraspecific competition may cause this behaviour. Future studies, which will test these possible explanations, will help to understand better the mechanism of co-evolutionary arms races and differences between host specialist and generalist brood parasites in various host–parasite systems.     

Rhythmic egg-laying behaviour in virgin females of fruit flies Drosophila melanogaster

Fruit fly Drosophila melanogaster females display rhythmic egg-laying under 12:12?h light/dark (LD) cycles which persists with near 24?h periodicity under constant darkness (DD). We have shown previously that persistence of this rhythm does not require the neurons expressing pigment dispersing factor (PDF), thought to be the canonical circadian pacemakers, and proposed that it could be controlled by peripheral clocks or regulated/triggered by the act of mating. We assayed egg-laying behaviour of wild-type Canton S (CS) females under LD, DD and constant light (LL) conditions in three different physiological states; as virgins, as females allowed to mate with males for 1?day and as females allowed to mate for the entire duration of the assay. Here, we report the presence of a circadian rhythm in egg-laying in virgin D. melanogaster females. We also found that egg-laying behaviour of 70 and 90% females from all the three male presence/absence protocols follows circadian rhythmicity under DD and LL, with periods ranging between 18 and 30?h. The egg-laying rhythm of all virgin females synchronized to LD cycles with a peak occurring soon after lights-off. The rhythm in virgins was remarkably robust with maximum number of eggs deposited immediately after lights-off in contrast to mated females which show higher egg-laying during the day. These results suggest that the egg-laying rhythm of D. melanogaster is endogenously driven and is neither regulated nor triggered by the act of mating; instead, the presence of males results in reduction in entrainment to LD cycles.     

Laying sequence,diploid males,and nest usurpation in the leafcutter bee,Megachile rotundata (Hymenoptera: Megachilidae)

Variations from the normal female-male sequence of eggs in nests of the leafcutter bee,Megachile rotundata, were examined. Three alternatives were considered: Out-of-sequence males (i) were diploids, (ii) were the result of supersedure of nests or intraspecific brood parasitism, or (iii) were the result of females occasionally laying a male-female sequence. Electrophoretic data provided definitive evidence of diploid males and of multiple females laying in 7 of 18 nests. In the others, the remaining explanation is that females occasionally lay male eggs before female eggs in a nest.     

Host nest usurpation and colony foundation in the European amazon ant,Polyergus rufescens Latr. (Hymenoptera: Formicidae)

Summary The socially parasitic mode of founding new colonies by queens of the European amazon antPolyergus rufescens was analysed in the laboratory. Newly-mated females of this obligatory slave-maker were individually introduced into queenright and queenless artificially established colonies of bothFormica cunicularia (the slave present in the natal dulotic nest) andF. rufibarbis (another potentialServiformica host). Particular attention was devoted to the behavioural patterns displayed by these young queens during the usurpation phases. Our observations, supported also by video-taping, show that the slave-making female, before laying her eggs, must penetrate the host colony, kill the resident queen, become accepted by the adult workers and appropriate the host brood. The parasite was almost always adopted in the colonies ofF. cunicularia, whereas in the presence ofF. rufibarbis it was generally killed in a short time. The failure in the attempt of usurping the colonies ofF. rufibarbis is discussed in relation to the host specificity typical of this slave-maker. Finally, egg-laying byPolyergus successful usurpers, the subsequent eclosion of the brood, and its complete social integration in the newly-established mixed colonies were also recorded.     

Apparent independence of the spawning and moulting cycles in female Antarctic krill (Euphausia superba Dana)

Summary Sixty female Antarctic krill (Euphausia superba Dana) spawned in shipboard experiments and the interval between egg-laying and ecdysis was noted. The number of eggs laid per female ranged from 263–3662, most females produced only one batch of eggs before moulting, and the post-spawn ovaries of all females contained few, if any, mature oocytes. As reported in other studies, the total number of eggs produced per female was not well correlated with body size. Females appeared to spawn at all times during the moulting cycle and although no diurnal rhythm in spawning was observed, moulting occurred mainly at night-time despite the animals being kept in near-constant darkness. No evidence of synchronous moutling was detected.     

Neuronal input contributes to sequence of Aplysia egg laying behaviors

Egg laying in Aplysia is controlled by the bag cell neuroendocrine system, which releases multiple peptides during a long-lasting electrical discharge. Following the discharge, a fixed sequence of head and neck movements is performed in which two phases can be distinguished: an appetitive or preparatory phase, in which the substrate is prepared, and a consummatory phase, when the egg string is deposited. During egg laying, feeding responses are suppressed. In this study, Aplysia fasciata was used. When the movement of the egg string through the genital groove was prevented by ligation, lesions of the nerve innervating the genital pore completely abolished the consummatory egg-laying behaviors. This shows that a nervous connection between the genital pore area and the central nervous system is important for the consummatory egg-laying behaviors.We found that suppression of feeding responses to seaweed occurred only during the consummatory phase of egg laying in controls, but animals with ligated genital grooves continued to show normal responses to food. It is hypothesized that a neuronal feedback, possibly together with the bag cell peptides, is critical for the temporal organization of egg-laying behavior in Aplysia.Abbreviations CNS central nervous system - ELH egg laying hormone     

The cost of reproduction in rosehip flies,Rhagoletis basiola: Eggs are time

Summary The hypothesis was tested that egg-laying by rosehip fruit flies,Rhagoletis basiola, would lead to reduced adult survivorship compared with oviposition-deprived flies, through experimental manipulation. Oviposition-deprived flies were assigned to two groups: mated and spinster. All three treatment groups were permitted to spend equal time and energy exploring and handling hosts but only the former group were allowed to deposit eggs (three/day). Results clearly showed that ovipositing flies suffered significantly higher mortality rates and that mated and spinster flies produced similar survivorship patterns. I consider these results both as a foraging problem (i.e. laying eggs in inferior hosts can cost both eggs and life) and as a more general life history problem.     

Shaking Youngsters and Shaken Adults: Female Beetles Eavesdrop on Larval Seed Vibrations to Make Egg-Laying Decisions

Egg-laying decisions are critical for insects, and particularly those competing for limited resources. Sensory information used by females to mediate egg-laying decisions has been reported to be primarily chemical, but the role of vibration has received little attention. We tested the hypothesis that vibrational cues produced by feeding larvae occupying a seed influences egg-laying decisions amongst female cowpea beetles. This hypothesis is supported by three lines of evidence using two strains of the cowpea beetle (Callosobruchus maculatus), an Indian strain with choosy females and aggressively competing larvae and a Brazilian strain with less choosy females and larvae exhibiting an “accommodating” type of competition. First, in free-choice bioassays of seed selection, choosy Indian females selected control seeds (free of eggs, larvae, or egg-laying marker) over seeds with live larvae (free of eggs and egg-laying marker), but did not discriminate between control seeds and those with dead larvae. In contrast, less choosy Brazilian females showed no preference for seeds containing live or dead larvae over controls. Second, laser-doppler vibrometer recordings confirmed that larvae feeding inside seeds generate vibrations that are available to the female during egg-laying decisions. Third, during dichotomous choice experiments where artificial vibrations approximating those produced by feeding larvae were played back during seed selection, Indian females preferred immobile control seeds over vibrating seeds, but Brazilian females showed no preference. These results support the hypothesis that females use larval vibrations in their egg-laying decisions; whether these vibrations are passive cues exploited by the female, or active signals that ‘steer’ the behaviour of the female is unknown. We propose that vibration cues and signals could be important for host selection in insects, particularly those laying on substrates where visual or chemical cues may be unreliable. This seems to be the case with females of the cowpea beetle since visual cues are not important and chemical egg-marking does not last more than two weeks, allowing vibration cues to improve discrimination of egg-laying substrate particularly by choosy females.     

The activity and distribution of wheat bulb fly (Delia coarctata) (Diptera:Anthomyiidae) in cereal crops and over fallow land

The flight activity and local distribution of adult D. coarctata were studied on Rothamsted Farm during 1970-5 using several trapping methods. Changes in the activity and distribution of the flies were followed as the population aged, and in females these changes were associated with maturation of their eggs. Most females appeared to stay at their emergence sites in winter wheat until egg-laying started about 1 month later. They then dispersed and both sexes were found in previously uninfested cereals and grass as well as on fallow land where the eggs are laid. Flight occurred chiefly in the late afternoon and evening. The concentration of flies at their emergence sites for several weeks before egg laying could facilitate chemical control, provided the practical difficulties of application can be overcome.     

Quantitative aspects of the effect of mating on readiness to lay in the Australian sheep blowfly,Lucilia cuprina

Virgin females of Lucilia cuprinararely lay eggs, whereas mated females do so readily. This effect of mating is due entirely to increased readiness to lay, and not to any effect on ovarian development. An investigation was made of how readiness to lay was affected by matings which differed in terms of the male's chemical and mechanical contribution. Individual males were mated, during 1 day, to a succession of females whose readiness to lay was determined 1 or 8 days after mating. On both days, the proportion of females laying was inversely related to the number of females with which the male had previously mated. A high proportion of females that had mated with previously unmated or oncemated males laid at both 1 and 8 days after mating. However, this proportion tended to decline between day 1 and day 8 in females that had mated with males with two or more previous matings, and this effect was most evident in females mated with males that had previously mated with four or more females. When matings were manually terminated as soon as coupling had occurred, the proportion laying remained as low as in virgins. This proportion progressively increased as mating duration increased from 2 to 6 min. The proportion that laid after mating terminated at 6 or 8 min was as high as that for females from full-term matings (mean duration, 12.5 min). The results are generally similar to those obtained in parallel experiments on the effect of mating on sexual receptivity in this species and, therefore, indicate that the physiological bases for the two effects of mating might be the same.     

Effects of carotenoid availability during laying on reproduction in the blue tit

Carotenoids are antioxidant pigments involved in several physiological processes and signalling in animals that cannot synthesise them and therefore must acquire them from food. We experimentally investigated the effects of carotenoid availability in the diet during egg laying on antioxidant deposition in egg yolk and the related effects on nestling condition, female body condition and parental investment in the blue tit (Parus caeruleus). Carotenoid supplementation of egg-laying females resulted in a significant increase in carotenoid concentration in egg yolk, but not in vitamin E or A concentration. There was no relationship between yellow plumage colour of adult females and carotenoid deposition in eggs, and no differential effect of feeding treatment depending on female colour. Nestlings from eggs laid by carotenoid supplemented females had longer tarsi, had faster development of the immune system as reflected by leukocyte concentration in blood, and grew brighter yellow feathers than nestlings from control females. However, nestlings from the two groups did not differ significantly in body mass, plasma antioxidants or plumage colour hue. At the time of chick rearing, carotenoid-fed females had increased plasma vitamin E levels compared to controls. However, females from the two treatment groups did not differ significantly in body condition or feeding rate. These results suggest that carotenoid availability is limiting during egg laying, and that females may have to balance the benefits of investing in egg quality against the potential costs of impairing their own future antioxidant protection. In addition, there may be considerable variation in carotenoid availability not only across seasons, but also among different stages of the breeding season.     

Experimentally induced clutch size enlargements affect reproductive success in the Pied Flycatcher

We performed a food provisioning experiment in a population of Pied Flycatchers Ficedula hypoleuca breeding at high altitude in central Spain to test if food availability before and during laying determines clutch size. Food was provided to one of two pairs with the same date of initiation of nest-building (15 dyads of subsequently reproducing pairs were thus created). Food provisioning began on the day of initiation of nest-building and ended on the day after the last egg was laid. Although laying date was unaffected by the experiment, clutch size in the experimental treatment was significantly larger. This result could indicate that food availability at laying (1) proximately constrained clutch size or (2) that females evaluated future conditions for incubating eggs and feeding nestlings based on food availability at laying. Reproductive success (proportion of eggs that resulted in fledged young) was significantly reduced in the experimental treatment. This effect suggest that supplemented females were tricked by the experiment into laying more eggs than the number of eggs they were able to incubate with success and the number of nestlings they were able to feed, a source of error in clutch size adjustment which could be common in non-experimental situations.