Oviposition by small hive beetles elicits hygienic responses from Cape honeybees

Two novel behaviours, both adaptations of small hive beetles (Aethina tumida Murray) and Cape honeybees (Apis mellifera capensis Esch.), are described. Beetles puncture the sides of empty cells and oviposit under the pupae in adjoining cells. However, bees detect this ruse and remove infested brood (hygienic behaviour), even under such well-disguised conditions. Indeed, bees removed 91% of treatment brood (brood cells with punctured walls caused by beetles) but only 2% of control brood (brood not exposed to beetles). Only 91% of treatment brood actually contained beetle eggs; the data therefore suggest that bees remove only that brood containing beetle eggs and leave uninfected brood alone, even if beetles have accessed (but not oviposited on) the brood. Although this unique oviposition strategy by beetles appears both elusive and adaptive, Cape honeybees are able to detect and remove virtually all of the infested brood.     

Explaining postnatal growth plasticity in a generalist brood parasite

Selection of a particular host has clear consequences for the performance of avian brood parasites. Experimental studies showed that growth rate and fledging mass of brood parasites varied between host species independently of the original host species. Finding correlates of this phenotypic plasticity in growth is important for assessing adaptiveness and potential fitness consequences of host choice. Here, I analyzed the effects of several host characteristics on growth rate and fledging mass of the young of brown-headed cowbird (Molothrus ater), a generalist, non-evicting brood parasite. Cowbird chicks grew better in fast-developing host species and reached higher fledging mass in large hosts with fast postnatal development. A potential proximate mechanism linking fast growth and high fledging mass of cowbird with fast host development is superior food supply in fast-developing foster species. So far, we know very little about the consequences of the great plasticity in cowbird growth for later performance of the adult parasite. Thus, cowbird species could become interesting model systems for investigating the role of plasticity and optimization in the evolution of growth rate in birds.     

Host genotype and age have no effect on rejection of parasitic eggs

Egg rejection belongs to a widely used host tactic to prevent the costs incurred by avian brood parasitism. However, the genetic basis of this behaviour and the effect of host age on the probability of rejecting the parasitic egg remain largely unknown. Here, we used a set of 15 polymorphic microsatellite loci, including a previously detected candidate locus (Ase64), to link genotypes of female great reed warblers (Acrocephalus arundinaceus), a known rejecter, with their egg rejection responses in two host populations. We also tested whether host female age, as a measure of the experience with own eggs, plays a role in rejection of common cuckoo (Cuculus canorus) eggs. We failed to find any consistent association of egg rejection responses with host female genotypes or age. It seems that host decisions on egg rejection show high levels of phenotypic plasticity and are likely to depend on the spatiotemporal variation in the parasitism pressure. Future studies exploring the repeatability of host responses towards parasitic eggs and the role of host individual experience with parasitic eggs would greatly improve our understanding of the variations in host behaviours considering the persistence of brood parasitism in host populations with rejecter phenotypes.     

Aggressive behavior of the male parent predicts brood sex ratio in a songbird

Brood sex ratio is often affected by parental or environmental quality, presumably in an adaptive manner that is the sex that confers higher fitness benefits to the mother is overproduced. So far, studies on the role of parental quality have focused on parental morphology and attractiveness. However, another aspect, the partner’s behavioral characteristics, may also be expected to play a role in brood sex ratio adjustment. To test this hypothesis, we investigated whether the proportion of sons in the brood is predicted by the level of territorial aggression displayed by the father, in the collared flycatcher (Ficedula albicollis). The proportion of sons in the brood was higher in early broods and increased with paternal tarsus length. When controlling for breeding date and body size, we found a higher proportion of sons in the brood of less aggressive fathers. Male nestlings are more sensitive to the rearing environment, and the behavior of courting males may often be used by females to assess their future parental activity. Therefore, adjusting brood sex ratio to the level of male aggression could be adaptive. Our results indicate that the behavior of the partner could indeed be a significant determinant in brood sex ratio adjustment, which should not be overlooked in future studies.     

Disc size regulation in the brood cell building behavior of leaf-cutter bee, <Emphasis Type="Italic">Megachile tsurugensis</Emphasis>

The leaf-cutter bee, Megachile tsurugensis, builds a brood cell in a preexisting tunnel with leaf discs that she cuts in decreasing sizes and assembles them like a Russian matryoshka doll. By experimentally manipulating the brood cell, it was investigated how she regulates the size of leaf discs that fit in the brood cell’s internal volume. When the internal volume was artificially increased by removing a bulk of leaf discs, she decreased the leaf disc size, although increasing it would have made the leaf disc more fitting in the increased internal volume. As a reverse manipulation, when the internal volume was decreased by inserting a group of inner layers of preassembled leaf discs to a brood cell, she decreased the leaf disc size, so that the leaf disc could fit in the decreased internal volume. These results suggest that she uses at least two different mechanisms to regulate the disc size: the use of some internal memory about the degree of building work accomplished in the first and of sensory feedback of dimensional information at the construction site in the second manipulation, respectively. It was concluded that a stigmergic mechanism, an immediate sensory feedback from the brood cell changed by the building work, alone cannot explain the details of the bee’s behavior particularly with respect to her initial response to the first manipulation. For a more complete explanation of the behavior exhibited by the solitary bee, two additional behavioral elements, reinforcement of building activity and processing of dimensional information, were discussed along with stigmergy.     

Pheromone-modulated behavioral suites influence colony growth in the honey bee (<Emphasis Type="Italic">Apis mellifera</Emphasis>)

The success of a species depends on its ability to assess its environment and to decide accordingly which behaviors are most appropriate. Many animal species, from bacteria to mammals, are able to communicate using interspecies chemicals called pheromones. In addition to exerting physiological effects on individuals, for social species, pheromones communicate group social structure. Communication of social structure is important to social insects for the allocation of its working members into coordinated suites of behaviors. We tested effects of long-term treatment with brood pheromone on suites of honey bee brood rearing and foraging behaviors. Pheromone-treated colonies reared significantly greater brood areas and more adults than controls, while amounts of stored pollen and honey remained statistically similar. Brood pheromone increased the number of pollen foragers and the pollen load weights they returned. It appeared that the pheromone-induced increase in pollen intake was directly canalized into more brood rearing. A two-way pheromone priming effect was observed, such that some workers from the same age cohorts showed an increased and extended capacity to rear larvae, while others were recruited at significantly younger ages into pollen-specific foraging. Brood pheromone affected suites of nursing and foraging behaviors allocating worker and pollen resources associated with an important fitness trait, colony growth.     

Behavioral genomics of honeybee foraging and nest defense

The honeybee has been the most important insect species for study of social behavior. The recently released draft genomic sequence for the bee will accelerate honeybee behavioral genetics. Although we lack sufficient tools to manipulate this genome easily, quantitative trait loci (QTLs) that influence natural variation in behavior have been identified and tested for their effects on correlated behavioral traits. We review what is known about the genetics and physiology of two behavioral traits in honeybees, foraging specialization (pollen versus nectar), and defensive behavior, and present evidence that map-based cloning of genes is more feasible in the bee than in other metazoans. We also present bioinformatic analyses of candidate genes within QTL confidence intervals (CIs). The high recombination rate of the bee made it possible to narrow the search to regions containing only 17–61 predicted peptides for each QTL, although CIs covered large genetic distances. Knowledge of correlated behavioral traits, comparative bioinformatics, and expression assays facilitated evaluation of candidate genes. An overrepresentation of genes involved in ovarian development and insulin-like signaling components within pollen foraging QTL regions suggests that an ancestral reproductive gene network was co-opted during the evolution of foraging specialization. The major QTL influencing defensive/aggressive behavior contains orthologs of genes involved in central nervous system activity and neurogenesis. Candidates at the other two defensive-behavior QTLs include modulators of sensory signaling (Am5HT ₇ serotonin receptor, AmArr4 arrestin, and GABA-B-R1 receptor). These studies are the first step in linking natural variation in honeybee social behavior to the identification of underlying genes.     

Heat Shielding: A Novel Method of Colonial Thermoregulation in Honey Bees

 Honey bees, Apis mellifera, maintain constant colony temperatures throughout the year. Honey bees fan their wings to cool the colony, and often spread fluid in conjunction with this behavior to induce evaporative cooling. We present an additional, previously undescribed mechanism used by the honey bee to maintain constant colony temperature in response to localized temperature increases. Worker bees shield the comb from external heat sources by positioning themselves on hot interior regions of the hive's walls. Although honey comb and brood comb were both shielded, the temperature-sensitive brood received a greater number of heat shielders and was thus better protected from overheating. Heat shielding appears to be a context-dependent adaptive behavior performed by worker bees who would previously have been considered "unemployed." Received: 16 November 1998 / Accepted in revised form: 31 March 1999     

Mandible shape in hybrid mice

Hybridisation between closely related species is frequently seen as retarding evolutionary divergence and can also promote it by creating novel phenotypes due to new genetic combinations and developmental interactions. We therefore investigated how hybridisation affects the shape of the mouse mandible, a well-known feature in evo–devo studies. Parental groups corresponded to two strains of the European mouse sub-species Mus musculus domesticus and Mus musculus musculus. Parents and hybrids were bred in controlled conditions. The mandibles of F₁ hybrids are mostly intermediate between parental phenotypes as expected for a complex multigenic character. Nevertheless, a transgressive effect as well as an increased phenotypic variance characterise the hybrids. This suggests that hybridisation between the two subspecies could lead to a higher phenotypic variance due to complex interactions among the parental genomes including non-additive genetic effects. The major direction of variance is conserved, however, among hybrids and parent groups. Hybridisation may thus play a role in the production of original transgressive phenotypes occurring following pre-existing patterns of variance.     

Impact of environmental temperatures on mortality,sex and caste ratios in <Emphasis Type="Italic">Melipona interrupta</Emphasis> Latreille (Hymenoptera,Apidae)

Temperature is a major driver of biological phenomena, from metabolism to ecological interactions and rates of evolutionary diversification. However, species vary greatly in their thermal tolerance, as well as the temperature under which they perform best. This study aimed to investigate the effect of experimental manipulation of environmental temperatures on the individual mortality and phenotypic composition of colonies of Melipona interrupta. To fulfill these objectives, 30 colonies in equivalent developmental conditions were artificially subjected to different temperatures. Temperatures were monitored by thermo-hygrometers, and immature mortality and sex and caste ratios were observed in brood combs during 14 months. A strong effect of external temperature on immatures was detected on deviations from 28 to 30 °C (the natural average temperature inside the colony), causing an increase in mortality. Likewise, a significant effect of temperature on sex ratio was detected, with male:female ratio decreasing at temperatures below and above 28–30 °C. Lastly, there was no clear evidence for an effect of temperature on caste ratio, although queens appeared to become relatively more frequent at warmer temperatures. The results of this study allow us to conclude that anthropogenic changes, whose effect can be extrapolated to the similar natural changes, that modify the environmental temperatures to which M. interrupta colonies are exposed are likely to compromise their survival, mainly through individual mortality.     

Caps and gaps: a computer model for studies on brood incubation strategies in honeybees (<Emphasis Type="Italic">Apis mellifera carnica</Emphasis>)

In addition to heat production on the comb surface, honeybee workers frequently visit open cells (“gaps”) that are scattered throughout the sealed brood area, and enter them to incubate adjacent brood cells. We examined the efficiency of this heating strategy under different environmental conditions and for gap proportions from 0 to 50%. For gap proportions from 4 to 10%, which are common to healthy colonies, we find a significant reduction in the incubation time per brood cell to maintain the correct temperature. The savings make up 18 to 37% of the time, which would be required for this task in completely sealed brood areas without any gaps. For unnatural high proportions of gaps (>20%), which may be the result of inbreeding or indicate a poor condition of the colony, brood nest thermoregulation becomes less efficient, and the incubation time per brood cell has to increase to maintain breeding temperature. Although the presence of gaps is not essential to maintain an optimal brood nest temperature, a small number of gaps make heating more economical by reducing the time and energy that must be spent on this vital task. As the benefit depends on the availability, spatial distribution and usage of gaps by the bees, further studies need to show the extent to which these results apply to real colonies. M. Fehler and M. Kleinhenz contributed equally to this work.     

基于逆方差多模型融合的空气质量指数预测方法

空气质量预测对合理制定环境治理政策具有重要意义。针对目前单体预测模型存在模型不稳定和泛化能力不强的问题,提出基于逆方差权重分配方法融合3种单体模型的空气质量指数(air quality index, AQI)预测方法。首先,以北京市为例,构建空气质量指数预测数据集;其次,分别构建长短期记忆网络(LSTM)、门控循环单元(GRU)、双向长短期记忆网络(Bi-LSTM)、自回归积分滑动平均模型(ARIMA)和多元线性回归(MLR)5种模型对数据集进行预测,并对比以上模型的预测结果;最后,在多模型融合方法中,选择逆方差法计算预测精度较高的3种单体模型的权重,根据算得权重构建逆方差融合预测模型。与预测精度较高的3种单体模型以及加权平均融合预测模型相比,逆方差融合预测模型对空气质量指数的预测精度R²分别提高3.9%、3.4%、1.6%和0.5%,达到0.933。结果表明：逆方差融合预测模型综合了各单体预测模型的优点,能够提高AQI预测精度。     

Colony size affects division of labour in the ponerine ant<Emphasis Type="Italic"> Rhytidoponera metallica</Emphasis>

In theory, larger colonies of social insects should have greater colony organisation. While inter-specific comparative studies provide support for this idea, there is little direct intra-specific evidence. We investigated differences in task specialisation between large (>450 workers) and small (<80 workers) colonies of the ponerine ant Rhytidoponera metallica. Observations of individually marked young or old workers revealed greater task specialisation in large colonies. Age polyethism was detected in large but not small colonies. In large colonies, old workers spent significantly more time foraging than young workers did, while young workers spent more time caring for brood. In small colonies, young and old workers spent a similar amount of time foraging and caring for brood. This difference in task allocation patterns in large and small colonies was associated with a difference in contact rates between workers. Workers in small colonies have a lower contact rate between nestmates and a greater variability in time between contacts than workers from large colonies.     

Hot spots in the bee hive

Honeybee colonies (Apis mellifera) maintain temperatures of 35-36°C in their brood nest because the brood needs high and constant temperature conditions for optimal development. We show that incubation of the brood at the level of individual honeybees is done by worker bees performing a particular and not yet specified behaviour: such bees raise the brood temperature by pressing their warm thoraces firmly onto caps under which the pupae develop. The bees stay motionless in a characteristic posture and have significantly higher thoracic temperatures than bees not assuming this posture in the brood area. The surface of the brood caps against which warm bees had pressed their thorax were up to 3.2°C warmer than the surrounding area, confirming that effective thermal transfer had taken place.     

Predatory behavior in a necrophagous bee<Emphasis Type="Italic"> Trigona hypogea</Emphasis> (Hymenoptera; Apidae,Meliponini)

Although most bees feed on nectar and pollen, several exceptions have been reported. The strangest of all is the habit found in some neotropical stingless bees, which have completely replaced pollen-eating by eating animal protein from corpses. For more than 20 years, it was believed that carrion was the only protein source for these bees. We report that these bees feed not only off dead animals, but on the living brood of social wasps and possibly other similar sources. Using well developed prey location and foraging behaviors, necrophagous bees discover recently abandoned wasps nests and, within a few hours, prey upon all immatures found there.     

Evidence of cue synergism in termite corpse response behavior

Subterranean termites of the genus Reticulitermes are known to build walls and tubes and move considerable amounts of soil into wood but the causes of this behavior remain largely unexplored. In laboratory assays, we tested the hypothesis that Reticulitermes virginicus (Banks) would carry more sand into wooden blocks containing corpses compared to corpse-free controls. We further predicted that the corpses of predatory ants would elicit a stronger response than those of a benign beetle species or nestmates. As hypothesized, significantly more sand was carried into blocks containing corpses and this material was typically used to build partitions separating the dead from the rest of the colony. Contrary to expectations, however, this behavior did not vary among corpse types. We then tested the hypothesis that oleic acid, an unsaturated fatty acid released during arthropod decay and used by ants and other arthropod taxa in corpse recognition, would induce a similar building response in R. virginicus. To additionally determine the role of foreign objects in giving rise to this behavior, the experiment was carried out with and without imitation corpses (i.e., small glass beads). As predicted, oleic acid induced building (a tenfold increase) but only when applied to beads, suggesting strong synergism between tactile and chemical cues. Oleic acid also significantly reduced the amount of wood consumed by R. virginicus and may possess useful repellent properties.     

Keeping eggs warm: thermal and developmental advantages for parasitic cuckoos of laying unusually thick-shelled eggs

Obligate brood parasites have evolved unusually thick-shelled eggs, which are hypothesized to possess a variety of functions such as resistance to puncture ejection by their hosts. In this study, we tested the hypothesis that obligate brood parasites lay unusually thick-shelled eggs to retain more heat for the developing embryo and thus contribute to early hatching of parasite eggs. By doing so, we used an infrared thermal imaging system as a non-invasive method to quantify the temperature of eggshells of common cuckoos (Cuculus canorus) and their Oriental reed warbler (Acrocephalus orientalis) hosts in an experiment that artificially altered the duration of incubation. Our results showed that cuckoo eggshells had higher temperature than host eggs during incubation, but also less fluctuations in temperature during incubation disturbance. Therefore, there was a thermal and hence a developmental advantage for brood parasitic cuckoos of laying thick-shelled eggs, providing another possible explanation for the unusually thick-shelled eggs of obligate brood parasites and earlier hatching of cuckoo eggs compared to those of the host.     

Plant intelligence

Intelligent behavior is a complex adaptive phenomenon that has evolved to enable organisms to deal with variable environmental circumstances. Maximizing fitness requires skill in foraging for necessary resources (food) in competitive circumstances and is probably the activity in which intelligent behavior is most easily seen. Biologists suggest that intelligence encompasses the characteristics of detailed sensory perception, information processing, learning, memory, choice, optimisation of resource sequestration with minimal outlay, self-recognition, and foresight by predictive modeling. All these properties are concerned with a capacity for problem solving in recurrent and novel situations. Here I review the evidence that individual plant species exhibit all of these intelligent behavioral capabilities but do so through phenotypic plasticity, not movement. Furthermore it is in the competitive foraging for resources that most of these intelligent attributes have been detected. Plants should therefore be regarded as prototypical intelligent organisms, a concept that has considerable consequences for investigations of whole plant communication, computation and signal transduction.     

Does the Spatial Distribution of the Parasitic Mite Varroa jacobsoni Oud. (Mesostigmata: Varroidae) in Worker Brood of Honey Bee Apis Mellifera L. (Hymenoptera: Apidae) Rely on an Aggregative Process?

  Varroa jacobsoni is an ectoparasite of honey bees which reproduces in capped brood cells. Multi-infestation is frequently observed in worker brood and can be interpreted as an aggregative phenomenon. The aim of this study was to determine whether the distribution of V. jacobsoni in worker brood cells relies on a random or an aggregative process. We studied the distribution of Varroa females in capped worker brood at similar age by comparing, by a Monte Carlo test, the observed frequency distribution of mites per cell to simulated distributions based on a random process. A complementary approach, using the "nearest neighbor distances" (NND) with Monte Carlo tests, was investigated to study the spatial distribution (a) between mites in different cells and (b) between infested cells in brood. The observed distributions did not differ significantly from that expected by a random process, and we conclude that there is no aggregation during invasion of V. jacobsoni in worker brood. Received: 29 April 1999 / Accepted in revised form: 26 August 1999