Non-transferable signals on ant queen eggs

How biological systems resolve internal conflicts is a major evolutionary question. Social insect workers cooperate but also pursue individual interests, such as laying male eggs. The rewards of this individual selfishness can be reduced by policing, such as by killing worker-laid eggs. However, selfish individuals may evade policing. What factors prevent individuals from being able to evade policing? In the ant Pachycondyla inversa, workers kill (police) worker-laid eggs. Because the colony keeps eggs in piles and worker-laid and queen-laid eggs are chemically distinct, worker-laid eggs might become more acceptable once placed in the egg pile by odour transfer from touching queen-laid eggs. Here, we show that such “cue scrambling” does not occur. Worker-laid eggs that were sandwiched between three queen-laid eggs for 45 min were not more acceptable in a policing bioassay than control worker-laid eggs. Chemical analyses also showed that the surface hydrocarbon profile of these eggs was unchanged. Policing, therefore, is stable against this potential cheating mechanism probably because queen-laid eggs are made chemically distinct using chemicals, that are not easily transferred by physical contact.     

A non-policing honey bee colony (Apis mellifera capensis)

In the Cape honey bee Apis mellifera capensis, workers lay female eggs without mating by thelytokous parthenogenesis. As a result, workers are as related to worker-laid eggs as they are to queen-laid eggs and therefore worker policing is expected to be lower, or even absent. This was tested by transferring worker- and queen-laid eggs into three queenright A. m. capensis discriminator colonies and monitoring their removal. Our results show that worker policing is variable in A. m. capensis and that in one colony worker-laid eggs were not removed. This is the first report of a non-policing queenright honey bee colony. DNA microsatellite and morphometric analysis suggests that the racial composition of the three discriminator colonies was different. The variation in policing rates could be explained by differences in degrees of hybridisation between A. m. capensis and A. m. scutellata, although a larger survey is needed to confirm this.     

Surface lipids of queen-laid eggs do not regulate queen production in a fission-performing ant

In animal societies, most collective and individual decision making depends on the presence of reproductive individuals. The efficient transmission of information among reproductive and non-reproductive individuals is therefore a determinant of colony organization. In social insects, the presence of a queen modulates multiple colonial activities. In many species, it negatively affects worker reproduction and the development of diploid larvae into future queens. The queen mostly signals her presence through pheromone emission, but the means by which these chemicals are distributed in the colony are still unclear. In several ant species, queen-laid eggs are the vehicle of the queen signal. The aim of this study was to investigate whether queen-laid eggs of the ant Aphaenogaster senilis possess queen-specific cuticular hydrocarbons and/or Dufour or poison gland compounds, and whether the presence of eggs inhibited larval development into queens. Our results show that the queen- and worker-laid eggs shared cuticular and Dufour hydrocarbons with the adults; however, their poison gland compounds were not similar. Queen-laid eggs had more dimethylalkanes and possessed a queen-specific mixture of cuticular hydrocarbons composed of 3,11?+?3,9?+?3,7-dimethylnonacosane, in higher proportions than did worker-laid eggs. Even though the queen-laid eggs were biochemically similar to the queen, their addition to experimentally queenless groups did not prevent the development of new queens. More studies are needed on the means by which queen ant pheromones are transmitted in the colony, and how these mechanisms correlates with life history traits.     

Mimicry of queen Dufour's gland secretions by workers of Apis mellifera scutellata and A. m. capensis

The development of the Dufour's gland of workers of the two honey bee races Apis mellifera scutellata and A. m. capensis was measured. The Dufour's glands of A. m. capensis workers were longer and increased in length more rapidly than the glands of workers of A. m. scutellata at comparable ages. Analysis of the Dufour's gland secretions of workers and queens of both races revealed that there were caste and racial differences. Secretions of queenright A. m. scutellata workers were dominated by a series of long-chain hydrocarbons. In contrast the secretions of the A. m. capensis workers both under queenright and queenless conditions were a mixture of hydrocarbons and wax-type esters, as were those of queens. Multivariate analysis of the secretion profiles indicated that laying workers of both races mimic queens. The secretions of the A. m. capensis laying workers mimicked queen secretions most closely, enabling them to act as successful social parasites.     

Similar policing rates of eggs laid by virgin and mated honey-bee queens

Worker-policing is a well-documented mechanism that maintains functional worker sterility in queenright honey-bee colonies. Unknown, however, is the source of the egg-marking signal that is thought to be produced by the queen and used by policing workers to discriminate between queen- and worker-laid eggs. Here we investigate whether mating is necessary for the queen to produce the egg-marking signal. We compare the removal rate of eggs laid by virgin queens and compare this rate with that of eggs laid by mated queens. Our results show that mating does not affect the acceptability of eggs, suggesting that physiological changes linked to the act of mating do not play a role in the production of the queens egg-marking signal.     

Can Dufour��s gland compounds honestly signal fertility in the primitively eusocial wasp Ropalidia marginata?

Unlike queens of typical primitively eusocial species, Ropalidia marginata queens are docile and non-interactive, and hence cannot be using dominance to maintain their status. It appears that the queen maintains reproductive monopoly through a pheromone, of which the Dufour's gland is at least one source. Here, we reconfirm earlier results showing that queens and workers can be correctly classified on a discriminant function using the compositions of their respective Dufour's glands, and also demonstrate consistent queen-worker differences based on categories of compounds and on single compounds also in some cases. Since the queen pheromone is expected to be an honest signal of the fecundity of a queen, we investigate the correlation of Dufour's gland compounds with ovarian activation of queens. Our study shows that Dufour's gland compounds in R. marginata correlate with the state of ovarian activation of queens, suggesting that such compounds may portray the fecundity of a queen, and may indeed function as honest signals of fertility.     

Nestmate recognition in social wasps: manipulation of hydrocarbon profiles induces aggression in the European hornet

The influence of individual cuticular hydrocarbons on nestmate recognition in the European hornet, Vespa crabro L., was investigated. We observed the behavioural response of workers towards differently treated dead conspecifics in a bioassay. Dummies were extracted with dichloromethane and extracts were spiked with microgram amounts of synthetic hydrocarbons naturally occurring on the cuticle of V. crabro. These modified extracts were reapplied to extracted workers that were subsequently tested in the bioassay. Non-spiked nestmate dummies (negative control) and untreated non-nestmate dummies (positive control) were tested in control experiments. The addition of only heneicosane or a mixture of heneicosane, tricosane, and (Z)-9-tricosene to the extracts led to a significant increase of agonistic behaviour in workers leaving the nest for foraging flights. Returning workers reacted much less aggressively than those leaving. This is one of the first behavioural proofs that manipulation of cuticular hydrocarbon profiles can be perceived by a social insect species. The results support the hypothesis that colony-specific cuticular hydrocarbon profiles are involved in the phenomenon of nestmate recognition among social insects.     

Experimental evidence that keeping eggs dry is a mechanism for the antimicrobial effects of avian incubation

Avian incubation dramatically reduces the abundance and diversity of microbial assemblages on eggshells, and this effect has been hypothesized as an adaptive explanation for partial incubation, the bouts of incubation that some birds perform during the egg-laying period. However, the mechanisms for these antimicrobial effects are largely unknown. In this study, we hypothesized that microbial inhibition is partly achieved through removal of liquid water, which generally enhances microbial growth, from eggshells, and experimentally tested this hypothesis in two ways. First, we placed the first- and second-laid eggs of tree swallow (Tachycineta bicolor) clutches in unincubated holding nests with either ambient or increased water on eggshells. Second, we added water to eggshells in naturally partially incubated nests. We compared microbial growth on shells during a 5-day experimental period and found that, as predicted, both unincubated groups had higher microbial growth than naturally partially incubated controls, and that only in the absence of incubation did wetted eggs have higher microbial growth than unwetted eggs. Thus, we have shown that water increases microbial growth on eggshells and that incubation nullifies these effects, suggesting that removal of water from egg surfaces is one proximate mechanism for the antimicrobial effects of incubation.     

Pre- and post-hatch trophic egg production in the subsocial burrower bug, Canthophorus niveimarginatus (Heteroptera: Cydnidae)

In recent years, three terrestrial bugs, Adomerus triguttulus and Sehirus cinctus (Cydnidae) and the closely related Parastrachia japonensis (Parastrachiidae), have been the focus of several fascinating studies because of the remarkable, extensive parental care they were found to display. This care includes egg and nymph guarding, production of trophic eggs, unfertilized, low cost eggs that are used as food by newly hatched nymphs, and progressive provisioning of the host seed. In this study, we have investigated yet a third related Asian cydnid, Canthophorus niveimarginatus, with regard to the possible occurrence of some or all of these complex traits in order to assess how widespread these maternal investment patterns are in this group of insects and to better understand the implications of their manifestations from an evolutionary context. Manipulative experiments were carried out in the lab to determine whether females provision nests. Observational and egg removal studies were carried out to determine whether trophic eggs are produced, and, if they are, their possible impact on nymphal success. The findings revealed that C. niveimarginatus does, in fact, progressively provision young, and this species also displays all of the other behaviors associated with extended parental care in subsocial insects. Moreover, unlike the other two related species, which place trophic eggs on the surface of the original egg mass, C. niveimarginatus produces both pre- and post-hatch trophic eggs. Nymphs deprived of access to post-hatch trophic eggs had significantly lower body weight and survival rate than those that fed on them. To our knowledge, this is the first time the production of both pre- and post-hatch trophic eggs has been demonstrated in insects outside the Hymenoptera. In this paper, we qualitatively and quantitatively demonstrate the provisioning behavior and patterns of trophic egg production in C. niveimarginatus. When and how trophic eggs are produced and delivered to young should have important correlations with the ecological and life history constraints under which a species has evolved. Thus, we also discuss the possible ecological and life history factors that favor the evolution of post-hatch trophic eggs.     

The role of tyramine and octopamine in the regulation of reproduction in queenless worker honeybees

In honeybees, workers under queenless condition compete for reproduction and establish reproductive dominance hierarchy. Ovary activation is generally accompanied by the expression of queen-like pheromones. Biogenic amines (BAs), in particular dopamine, are believed to be involved in this process by regulating ovarian development. However, the role of BAs in establishing reproductive dominance or their effect on queen-like pheromone production was not investigated. Here, we explored the effect of octopamine (OA) and tyramine (TA) oral treatments on the propensity of treated bees to become reproductively dominant and produce queen-like pheromones in Dufour’s and mandibular glands. One bee in a pair was treated with either OA or TA while the other was fed sugar solution. TA was found to enhance ovary development and the production of esters in the Dufour’s gland and 9HDA (queen component) in the mandibular glands, thus facilitating worker reproductive dominance. OA, on the other hand, did not enhance ovarian development or ester production, but increased the production of 10HDA (worker major component) in the mandibular glands of their sugar-paired mates. OA is known to induce foraging behavior by workers, while increased production of 10HDA characterizes nursing workers. Therefore, we suggest that TA induces reproductive division of labor, while OA treatment results in caste differentiation of workers to foragers and nurses.     

Brain modulation of Dufour’s gland ester biosynthesis in vitro in the honeybee (<Emphasis Type="Italic">Apis mellifera</Emphasis>)

Caste-specific pheromone biosynthesis is a prerequisite for reproductive skew in the honeybee. Nonetheless, this process is not hardwired but plastic, in that egg-laying workers produce a queen-like pheromone. Studies with Dufour’s gland pheromone revealed that, in vivo, workers’ gland biosynthesis matches the social status of the worker, i.e., sterile workers showed a worker-like pattern whereas fertile workers showed a queen-like pattern (production of the queen-specific esters). However, when incubated in vitro, the gland spontaneously exhibits the queen-like pattern, irrespective of its original worker type, prompting the notion that ester production in workers is under inhibitory control that is queen-dependent. We tested this hypothesis by exposing queen or worker Dufour’s glands in vitro to brain extracts of queens, queenright (sterile) workers and males. Unexpectedly, worker brain extracts activated the queen-like esters biosynthesis in workers’ Dufour’s gland. This stimulation was gender-specific; queen or worker brains demonstrated a stimulatory activity, but male brains did not. Queen gland could not be further stimulated. Bioassays with heated and filtered extracts indicate that the stimulatory brain factor is below 3,000 Da. We suggest that pheromone production in Dufour’s gland is under dual, negative–positive control. Under queenright conditions, the inhibitor is released and blocks ester biosynthesis, whereas under queenless conditions, the activator is released, activating ester biosynthesis in the gland. This is consistent with the hypothesis that queenright workers are unequivocally recognized as non-fertile, whereas queenless workers try to become “false queens” as part of the reproductive competition.     

Infanticide by a solitary koinobiont ichneumonid ectoparasitoid of spiders

When encountering an already parasitized host, a parasitoid’s optimal choices (superparasitism, host rejection, host feeding or infanticide) seem to depend on the individual species’ life history, because the same choice may have different fitness consequences. We demonstrate infanticide under laboratory conditions by a polysphinctine, Zatypota albicoxa, which is a solitary koinobiont ectoparasitoid of spiders. The female always removed any previously attached egg or larva from the body of the host spider, Parasteatoda tepidariorum, with a rubbing behaviour. She rubbed her ovipositor back and forth toward the undersurface of the attached egg or of the saddle under the attached larva to pry it off and laid an egg after removal. When removing a larva, the infanticidal female engaged exclusively in unfastening the ‘saddle’ which fastens the larva to the body of the spider. All larvae were removed with the ‘saddle’ attached to the ventral surface of the body. The female invested more time to remove the medium second and the large penultimate instar larvae than to remove eggs and first instar larvae because of the labour involved in unfastening the saddle. Oviposition with infanticide of the medium second and the penultimate instar larvae imposed more time upon the female than that on an unparasitized host. Removal of any previous occupant in spite of the associated labour costs suggests that infanticide will always be adaptive, no matter the time costs to Z. albicoxa, because so much is invested in attacking the host and because the parasitoid cannot detect whether the spider is already parasitized until she achieves subjugation.     

Hoopoes color their eggs with antimicrobial uropygial secretions

Uropygial gland secretions are used as cosmetics by some species of birds to color and enhance properties of feathers and teguments, which may signal individual quality. Uropygial secretions also reach eggshells during incubation and, therefore, may influence the coloration of birds’ eggs, a trait that has attracted the attention of evolutionary biologists for more than one century. The color of hoopoe eggs typically changes along incubation, from bluish-gray to greenish-brown. Here, we test experimentally the hypothesis that dark uropygial secretion of females is responsible for such drastic color change. Moreover, since uropygial secretion of hoopoes has antimicrobial properties, we also explore the association between color and antimicrobial activity of the uropygial secretion of females. We found that eggs stayed bluish-gray in nests where female access to the uropygial secretion was experimentally blocked. Furthermore, experimental eggs that were maintained in incubators and manually smeared with uropygial secretion experienced similar color changes that naturally incubated eggs did, while control eggs that were not in contact with the secretions did not experience such color changes. All these results strongly support the hypothesis that female hoopoes use their uropygial gland secretion to color the eggs. Moreover, saturation of the uropygial secretion was associated with antimicrobial activity against Bacillus licheniformis. Given the known antimicrobial potential of uropygial secretions of birds, this finding opens the possibility that in scenarios of sexual selection, hoopoes in particular and birds in general signal antimicrobial properties of their uropygial secretion by mean of changes in egg coloration along incubation.     

Body size phenotypes are heritable and mediate fecundity but not fitness in the lepidopteran frugivore Cydia pomonella

The inheritance and functional roles of quantitative traits are central concerns of evolutionary ecology. We report two sets of experiments that investigated the heritability and reproductive consequences of body size phenotypes in a globally distributed lepidopteran frugivore, Cydia pomonella (L.). In our first set of experiments, we tested the hypotheses that (1) body size is heritable and (2) parental body size mediates egg production and offspring survival. Midparent–offspring regression analyses revealed that body mass is highly heritable for females and moderately heritable for males. The contribution of fathers to estimates of additive genetic variance was slightly greater than for mothers. Egg production increased with mean parental size, but offspring survival rates were equivalent. Based on this result, we tested two additional hypotheses in a second set of experiments: (3) male size moderates female egg production and egg fertility and (4) egg production, egg fertility, and offspring survival rate are influenced by female mating opportunities. Females paired with large males produced more eggs and a higher proportion of fertile eggs than females paired with small males. Females with multiple mating opportunities produced more fertile eggs than females paired with a single male. However, egg production and offspring survival rates were unaffected by the number of mating opportunities. Our experiments demonstrate that body mass is heritable in C. pomonella and that size phenotypes may mediate fecundity but not fitness. We conclude that male size can influence egg production and fertility, but female mate choice also plays a role in determining egg fertility.     

Aphid egg protection by ants: a novel aspect of the mutualism between the tree-feeding aphid Stomaphis hirukawai and its attendant ant Lasius productus

Aphids often form mutualistic associations with ants, in which the aphids provide the ants with honeydew and the ants defend the aphids from predators. In this paper, we report aphid egg protection by ants as a novel aspect of the deeply interdependent relationship between a tree-feeding aphid and its attendant ant. The ant Lasius productus harbours oviparous females, males, and eggs of the hinoki cypress-feeding aphid Stomaphis hirukawai in its nests in winter. We investigated the behaviour of ants kept with aphid eggs in petri dishes to examine whether the ants recognise the aphid eggs and tend them or only provide a refuge for the aphids. Workers carried almost all of the aphid eggs into the nest within 24 h. The ants indiscriminately tended aphid eggs collected from their own colonies and those from other ant colonies. The ants cleaned the eggs and piled them up in the nest, and egg tending by ants dramatically increased aphid egg survival rates. Starving the ants showed no significant effect on aphid egg survivorship. Without ants, aphid eggs were rapidly killed by fungi. These results suggested that grooming by the ants protected the aphid eggs, at least, against pathogenic fungi. This hygienic service afforded by the ants seems indispensable for egg survival of these aphids in an environment rich in potentially pathogenic microorganisms.Electronic supplementary material  Supplementary material is available for this article at and is accessible for authorized users.     

纳滤膜去除卡马西平的影响因素研究

主要研究了纳滤膜去除饮用水中卡马西平(CBZ)的影响因素.首先比较了膜NF270和NF90去除CBZ的效果,发现孔径较小的NF90对CBZ的去除效果远优于孔径较大的NF270.其次考察了CBZ初始浓度、pH值、离子强度和水温对去除率的影响.结果表明,pH值的降低,钙离子浓度的提高以及水温的提高会导致CBZ去除率的降低....     

环境激素氯氰菊酯对萼花臂尾轮虫繁殖的影响

以2 d种群增长率为指标研究了环境抗雄激素氯氰菊酯对萼花臂尾轮虫的急性毒性;以3 d种群动态参数(种群增长率、混交率、混交雌体受精率和携卵雌体/非携卵雌体)、7 d休眠卵产量和休眠卵孵化率为指标研究了低剂量氯氰菊酯(0.001～0.316 mg.L-1)对萼花臂尾轮虫繁殖的影响;以2 d种群参数分析了在氯氰菊酯中形成休眠卵孵化后的生长性能;以3 d种群参数为指标分析了母体暴露氯氰菊酯,对其后代繁殖的影响.结果表明氯氰菊酯浓度对数与种群增长率呈直线负相关.氯氰菊酯半数效应浓度(EC50)、最低可观察效应浓度(LOEC)和无可观察效应浓度(NOEC)分别为14.22、10和3.16mg.L-1;0.0316 mg.L-1氯氰菊酯组7 d休眠卵产量较对照组下降了41.23%,休眠卵孵化率亦较对照组显著下降;氯氰菊酯试验组中形成的休眠卵孵化后的种群增长率和混交率显著下降;萼花臂尾轮虫母体暴露0.316 mg.L-1氯氰菊酯其后代种群增长率比对照下降了15.96%.试验表明,萼花臂尾轮虫2 d种群增长率对氯氰菊酯较不敏感;低剂量氯氰菊酯可降低休眠卵产量、休眠卵孵化率及孵化后种群增长率,从而减少萼花臂尾轮虫后代早期对种群增长的贡献.     

改性沉积物除藻对水质改善的效果研究

通过生态模拟箱(1m×1m×1m)模拟天然湖泊环境,研究了改性沉积物的除藻效果和对水质的改善作用.改性沉积物除藻后,测定和分析了除藻箱体和不除藻箱体内的叶绿素、浊度、DO、TN、TP、DIN、DIP等水质指标.在改性沉积物投量为43mg·L-1时,4h除藻率可达到85%以上.除藻2d后,水体中叶绿素和浊度降低90%,总氮、总磷分别降低70%、80%,这些指标在15d的连续监测期间变化不大.除藻后2箱体中DO都有不同程度的下降,但除藻箱体中DO明显高于不除藻箱体.并且在此实验期间,除藻箱体DIN降低,DIP稳定在较低水平,而不除藻箱体中DIN与DIP均有明显升高.说明改性沉积物除藻能有效减缓水体的厌氧情况,在短期内能抑制营养盐的释放.以上结果表明了改性沉积物在除藻的同时能有效改善水质,可用于水源地的水质保护.     

中试规模微气泡曝气生物膜反应器运行性能评估

运行中试规模微气泡曝气生物膜反应器处理校园生活污水,对其运行性能进行评估,并与传统生物处理工艺比较.结果表明,采用中试系统处理校园生活污水原水时,平均COD去除率和去除负荷分别为57.0%和2.68 kg·(m~3·d)~(-1),平均氨氮去除率和去除负荷分别为17.4%和0.17 kg·(m~3·d)~(-1),平均TN去除率和去除负荷分别为15.8%和0.21 kg·(m~3·d)~(-1),平均氧利用率达到100%.采用中试系统处理可生化性较差的生物接触氧化池出水,平均COD去除率和去除负荷分别为46.0%和1.53 kg·(m~3·d)~(-1);平均氨氮去除率和去除负荷分别为17.1%和0.32 kg·(m~3·d)~(-1);平均TN去除率和去除负荷分别为14.1%和0.28 kg·(m~3·d)~(-1);平均氧利用率高于50%.由于微气泡曝气能够加速氧传质过程并提高氧利用率,因此相同进水条件下,中试系统污染物去除能力显著优于传统生物接触氧化工艺和传统曝气生物滤池工艺.     

模拟水泥窑工艺对污染土壤热解吸尾气中六氯苯的去除效果

为了降低HCB(六氯苯)污染土壤热解吸修复的成本,分析了水泥窑处理技术去除污染土壤热解吸尾气中HCB的可行性,主要考察了处理温度、停留时间、φ(O₂)和ρ(HCB)初始值对HCB去除的影响. 结果表明:①随着处理温度的升高和停留时间的延长,HCB的去除率逐渐升高,其中,当处理温度≥800 ℃、停留时间≥2 s时,其去除率高于99.93%. ②当处理温度为900 ℃、停留时间为2 s、ρ(HCB)初始值分别为1.70、17.00和85.00 mg/m3时,水泥窑处理后尾气中ρ(HCB)分别为0.60、0.78和1.50 μg/m3,其与ρ(HCB)初始值并不成正比,说明ρ(HCB)初始值对HCB去除的影响较小;φ(O₂)≥2%时对HCB去除的影响也较小. ③水泥窑处理后,HCB的脱氯降解产物中仅有五氯苯被检出,ρ(五氯苯)最大值为1.20 μg/m3;同时处理过程中伴有少量的二英产生,但ρ(二英)满足GB 30485—2013《水泥窑协同处置固体废物污染控制标准》要求. 结果显示,模拟水泥窑工艺可有效去除污染土壤热解吸尾气中的HCB.