A quantitative model of successive color induction in the honeybee

Intensity discrimination experiments are performed with individual walking honeybees trained to color stimuli (UV, blue and green) of constant intensity. The choice behavior to stimuli of identical wavelength spectrum but different intensities is tested. A graded choice behavior is found. The training intensity is chosen with the highest probability in most cases. Phototaxis as well as brightness discrimination can be excluded. The choice behavior is explained exclusively by discrimination of chromaticness (hue and saturation) according to the Bezold-Brücke shift.The bees adapt to the chromatic stimuli during their choices. From the behavioral data, it is concluded that in adaptation, adjustment in photoreceptor sensitivity in one receptor also affects the sensitivity of the other receptors (co-adaptation). The linear adaptation model corresponding to the von Kries Coefficient Law used up to now to describe adaptation to white light in the honeybee does not describe this type of adaptation.A quantitative model of adaptation to chromatic stimuli extending the linear adaptation model is developed.The most reasonable mechanism of co-adaptation is optical coupling by lateral filtering. Other mechanisms such as electrical coupling are unlikely, since their effects on color vision would lead to effects inconsistent with Graßmann's Laws.     

Chromatic and achromatic stimulus discrimination of long wavelength (red) visual stimuli by the honeybee Apis mellifera

It has long been assumed that bees cannot see red. However, bees visit red flowers, and the visual spectral sensitivity of bees extends into wavelengths to provide sensitivity to such flowers. We thus investigated whether bees can discriminate stimuli reflecting wavelengths above 560 nm, i.e., which appear orange and red to a human observer. Flowers do not reflect monochromatic (single wavelength) light; specifically orange and red flowers have reflectance patterns which are step functions, we thus used colored stimuli with such reflectance patterns. We first conditioned honey bees Apis mellifera to detect six stimuli reflecting light mostly above 560 nm and found that bees learned to detect only stimuli which were perceptually very different from a bee achromatic background. In a second experiment we conditioned bees to discriminate stimuli from a salient, negative (un-rewarded) yellow stimulus. In subsequent unrewarded tests we presented the bees with the trained situation and with five other tests in which the trained stimulus was presented against a novel one. We found that bees learned to discriminate the positive from the negative stimulus, and could unambiguously discriminate eight out of fifteen stimulus pairs. The performance of bees was positively correlated with differences between the trained and the novel stimulus in the receptor contrast for the long-wavelength bee photoreceptor and in the color distance (calculated using two models of the honeybee colors space). We found that the differential conditioning resulted in a concurrent inhibitory conditioning of the negative stimulus, which might have improved discrimination of stimuli which are perceptually similar. These results show that bees can detect long wavelength stimuli which appear reddish to a human observer. The mechanisms underlying discrimination of these stimuli are discussed. Handling Editor: Lars Chittka.     

Paternal effects on the defensive behavior of honeybees

The defensive behavior of 52 hybrid honeybee (Apis mellifera L.) colonies from four sets of crosses was studied and compared with that of European and Africanized bee colonies. Colonies containing F(1) hybrid workers were obtained through reciprocal crosses between European and Africanized bees. The total number of stings deposited by workers in a moving leather patch in 1 min was recorded. In each of the four sets of crosses, bees from hybrid colonies of Africanized paternity left more stings in leather patches than bees from hybrid colonies of European paternity. Results strongly suggest paternal effects of African origin increasing the defensive behavior of hybrid colonies. Although some degree of dominance was observed for high-defensive behavior in one of the four sets of crosses involving European paternity, most of the dominance effects reported in the literature appear to be the result of paternal effects. Several hypotheses to explain this phenomenon, as well as the implications of these effects on the fitness and breeding of honeybees are discussed.     

Imidacloprid-induced facilitation of the proboscis extension reflex habituation in the honeybee.

Imidacloprid is a new insecticide from the family of the neonicotinoids, which interact with the insect nicotinic acetylcholine receptor. The effects of imidacloprid at a dose that does not affect sensory or motor functions are studied on non-associative learning abilities in the honeybee. The behavioral procedure is the habituation of the proboscis extension reflex (PER). Imidacloprid topically applied on the thorax (1 microl) at the doses of 5, 10, and 20 ng/bee induces an increase of the gustatory threshold defined as the lowest concentration of a sucrose solution applied to the antennae able to elicit the PER. The ability of the honeybee to move in an open-field-like apparatus is impaired at the doses of 2.5, 5, 10, and 20 ng/bee. These effects are amplified with time and reach a maximum 60 min after application. The lowest dose of 1.25 ng/bee has no effect on the gustatory function but increases the motor activity and facilitates the PER habituation independently of time. This result fits our hypothesis that a slight activation of the cholinergic system with a low dose of imidacloprid can facilitate a simple form of learning in the honeybee.     

European honeybee defense against Japanese yellow hornet using heat generation by bee‐balling behavior

The Japanese honeybee, Apis cerana japonica Radoszkowski, uses unique generation of heat by bee‐balling to defend against, overheat and kill predacious Japanese hornets. We have now observed the European honeybee, Apis mellifera Linnaeus, using similar bee‐balling behavior and heat generation against the Japanese yellow hornet, Vespa simillima xanthoptera Cameron. We monitored temperatures in the center of the bee‐ball and inside thoracic muscles of the captured hornet and found that the thoracic internal temperature (45.8 ± 2.32°C) was higher than that of the bee‐ball (44.0 ± 0.96°C). Although the thoracic temperature of captured hornets rose to the upper lethal level, defending European bees also showed some stinging attempts against the hornet, unlike the sympatric Japanese honeybee, which never stings during bee‐balling. The European honeybee bee‐balling behavior consists of three phases: (i) heating; (ii) heat‐retaining; and (iii) break up. Our results suggest that European honeybees kill hornets by raising the body temperature of hornets rapidly without stinging. The tactics of bee‐balling against hornets are complex and may be performed by extended division of labor.     

Successive two-item same-different discrimination and concept learning by pigeons

Four pigeons were trained in a successive same/different procedure involving the alternation of two stimuli per trial. Using a go/no-go procedure, two different or two identical color photographs were alternated, with a brief, dark, inter-stimulus interval, on a computer screen for 20s. Pigeons learned to discriminate between same (S+) and different (D-) sequences with moderate to large contrasts between successive pictures. Analyses of pecking behavior within single trials revealed this discrimination emerged at the earliest possible point in the sequence (i.e. by the presentation of the second item). Pigeons transferred to novel color and gray-scale pictures, and showed savings in tests with novel video stimuli. These results suggest that same/different discrimination and concept formation can be acquired with successively presented pairs of stimuli by pigeons. When combined with results using simultaneous same/different presentations, these findings further support a qualitative similarity among birds and primates in their capacity to judge certain types of stimulus relations.     

An investigation of the role of dopamine in the antennal lobes of the honeybee,Apis mellifera

Summary The effects of dopamine applied to the antennal lobes of the honeybee are investigated using the proboscis conditioning paradigm (Kuwabara 1957). The percentage of bees responding to a conditioned olfactory stimulus after a single conditioning trial is reduced significantly by the application of dopamine (10^–6 M) to the antennal lobes of the bee brain. Reduction in response levels is significantly smaller in bees conditioned to the olfactory stimulus in multiple conditioning trials prior to treatment with dopamine. The effects of dopamine on the percentage of bees responding to a conditioned olfactory stimulus are blocked by the butyrophenone, haloperidol (10^–5 M). The possible role of dopaminergic interneurones in the antennal lobes of the bee brain is discussed.     

The Rat Electroretinogram : I. Contrasting effects of adaptation on the amplitude and latency of the b-wave

Characteristics of the electroretinogram (ERG) produced by the essentially all rod eye of the rat are presented as functions of the number of quanta absorbed by each rod per stimulus flash. The ERG's were obtained with 1.5 msec. stimulus flashes and uniform illumination of the entire retina. Under these conditions, distortions in the ERG due to stray light are minimized, and the ERG more accurately reflects the activity of its retinal sources. The effects of background light and two forms of dark adaptation were studied and compared. The results, especially for the b-wave, permit an interpretation in terms of two distinct processes. One process appears to determine the b-wave latency. This process is almost independent of the state of adaptation of the retina. The other process does not affect the latency, but determines the b-wave threshold and amplitude. This process strongly depends upon the state of adaptation. Moreover, the effects of dark adaptation on this amplitude-determining process are almost identical with the effects of background light.     

RFID tracking of sublethal effects of two neonicotinoid insecticides on the foraging behavior of Apis mellifera

The development of insecticides requires valid risk assessment procedures to avoid causing harm to beneficial insects and especially to pollinators such as the honeybee Apis mellifera. In addition to testing according to current guidelines designed to detect bee mortality, tests are needed to determine possible sublethal effects interfering with the animal's vitality and behavioral performance. Several methods have been used to detect sublethal effects of different insecticides under laboratory conditions using olfactory conditioning. Furthermore, studies have been conducted on the influence insecticides have on foraging activity and homing ability which require time-consuming visual observation. We tested an experimental design using the radiofrequency identification (RFID) method to monitor the influence of sublethal doses of insecticides on individual honeybee foragers on an automated basis. With electronic readers positioned at the hive entrance and at an artificial food source, we obtained quantifiable data on honeybee foraging behavior. This enabled us to efficiently retrieve detailed information on flight parameters. We compared several groups of bees, fed simultaneously with different dosages of a tested substance. With this experimental approach we monitored the acute effects of sublethal doses of the neonicotinoids imidacloprid (0.15-6 ng/bee) and clothianidin (0.05-2 ng/bee) under field-like circumstances. At field-relevant doses for nectar and pollen no adverse effects were observed for either substance. Both substances led to a significant reduction of foraging activity and to longer foraging flights at doses of ≥0.5 ng/bee (clothianidin) and ≥1.5 ng/bee (imidacloprid) during the first three hours after treatment. This study demonstrates that the RFID-method is an effective way to record short-term alterations in foraging activity after insecticides have been administered once, orally, to individual bees. We contribute further information on the understanding of how honeybees are affected by sublethal doses of insecticides.     

Asynchronous processing in vision: color leads motion

It has been demonstrated that subjects do not report changes in color and direction of motion as being co-incidental when they occur synchronously. Instead, for the changes to be reported as being synchronous, changes in direction of motion must precede changes in color. To explain this observation, some researchers have suggested that the neural processing of color and motion is asynchronous. This interpretation has been criticized on the basis that processing time may not correlate directly and invariantly with perceived time of occurrence. Here we examine this possibility by making use of the color-contingent motion aftereffect. By correlating color states disproportionately with two directions of motion, we produced and measured color-contingent motion aftereffects as a function of the range of physical correlations. The aftereffects observed are consistent with the perceptual correlation between color and motion being different from the physical correlation. These findings demonstrate asynchronous processing for different stimulus attributes, with color being processed more quickly than motion. This suggests that the time course of perceptual experience correlates directly with that of neural activity.     

Neural adjustments to chromatic blur

The perception of blur in images can be strongly affected by prior adaptation to blurry images or by spatial induction from blurred surrounds. These contextual effects may play a role in calibrating visual responses for the spatial structure of luminance variations in images. We asked whether similar adjustments might also calibrate the visual system for spatial variations in color. Observers adjusted the amplitude spectra of luminance or chromatic images until they appeared correctly focused, and repeated these measurements either before or after adaptation to blurred or sharpened images or in the presence of blurred or sharpened surrounds. Prior adaptation induced large and distinct changes in perceived focus for both luminance and chromatic patterns, suggesting that luminance and chromatic mechanisms are both able to adjust to changes in the level of blur. However, judgments of focus were more variable for color, and unlike luminance there was little effect of surrounding spatial context on perceived blur. In additional measurements we explored the effects of adaptation on threshold contrast sensitivity for luminance and color. Adaptation to filtered noise with a 1/f spectrum characteristic of natural images strongly and selectively elevated thresholds at low spatial frequencies for both luminance and color, thus transforming the chromatic contrast sensitivity function from lowpass to nearly bandpass. These threshold changes were found to reflect interactions between different spatial scales that bias sensitivity against the lowest spatial grain in the image, and may reflect adaptation to different stimulus attributes than the attributes underlying judgments of image focus. Our results suggest that spatial sensitivity for variations in color can be strongly shaped by adaptation to the spatial structure of the stimulus, but point to dissociations in these visual adjustments both between luminance and color and different measures of spatial sensitivity.     

A Comparative Study of Relational Learning Capacity in Honeybees (Apis mellifera) and Stingless Bees (Melipona rufiventris)

Background

Learning of arbitrary relations is the capacity to acquire knowledge about associations between events or stimuli that do not share any similarities, and use this knowledge to make behavioural choices. This capacity is well documented in humans and vertebrates, and there is some evidence it exists in the honeybee (Apis mellifera). However, little is known about whether the ability for relational learning extends to other invertebrates, although many insects have been shown to possess excellent learning capacities in spite of their small brains.

Methodology/Principal Findings

Using a symbolic matching-to-sample procedure, we show that the honeybee Apis mellifera rapidly learns arbitrary relations between colours and patterns, reaching 68.2% correct choice for pattern-colour relations and 73.3% for colour-pattern relations. However, Apis mellifera does not transfer this knowledge to the symmetrical relations when the stimulus order is reversed. A second bee species, the stingless bee Melipona rufiventris from Brazil, seems unable to learn the same arbitrary relations between colours and patterns, although it exhibits excellent discrimination learning.

Conclusions/Significance

Our results confirm that the capacity for learning arbitrary relations is not limited to vertebrates, but even insects with small brains can perform this learning task. Interestingly, it seems to be a species-specific ability. The disparity in relational learning performance between the two bee species we tested may be linked to their specific foraging and recruitment strategies, which evolved in adaptation to different environments.     

Phenotypic plasticity in insects: the effects of substrate color on the coloration of two ground-hopper species

SUMMARY The question of how phenotypic variation is maintained within populations has long been a central issue in evolutionary biology. Most of these studies focused on the maintenance of genetic variability, but the phenotype of organisms may also be influenced by environmental cues experienced during ontogeny. Color polymorphism has received particular attention in evolutionary studies as it has strong fitness consequences. However, if body coloration is influenced by the environment, any conclusions on evolutionary consequences of fitness trade-offs can be misleading. Here we present data from a laboratory experiment on the influence of substrate color on three aspects of the coloration of two ground-hopper species, Tetrix subulata and Tetrix ceperoi . We reared hatchlings either on dark or on light substrates, using a split-brood design. Although the type of pronotal pattern changed mainly in response to nymphal development, the basic color was strongly influenced by the substrate color. In both species, black and dark olive color morphs were found more frequently on the dark substrate, whereas the gray color morph dominated on the light substrate. These findings have considerable implications for our understanding of color morph evolution as they show that color polymorphism may not only be maintained by natural selection acting on discrete color morphs, but also by phenotypic plasticity, which enables organisms to adjust to the environmental conditions experienced during ontogeny. This facultative morphology is opposing to the prevailing view of color morph adaptation, which assumes a purely genetic determination and co-evolution of discrete color morphs with life history traits.     

Role of intracellular calcium and sodium in light adaptation in the retina of the honey bee drone (Apis mellifera, L)

In the honey bee drone, the decrease in sensitivity to light of a retinula cell exposed to background illumination was found to be accurately reflected by the difference in amplitude between the initial transient depolarization and the lowest steady depolarization evoked by the background light. It is shown that both the decrease in sensitivity to light and the accompanying drop in potential from the transient to the plateau can be prevented by injecting EGTA intracellularly. A decrease in duration and amplitude of responses to short test flashes such as observed immediately after illumination was found to occur too when Ca or Na, but not K, Li, or Mg injected into dark-adapted retinula cells. Injection of EGTA into a retinula cell maintained a steady state of light adaptation, was found to cause an increase in amplitude and duration of the response to a short test flash, thus producing the effects of dark adaptation. It is suggested that, in the retina of the honey bee drone, an increase in intracellular calcium concentration plays a central role in light adaptation and that an increase in intracellular sodium concentration, resulting from the influx of sodium ions during the responses to light, could lead to this increase in intracellular free calcium.     

Farming with native bees (<Emphasis Type="Italic">Apis mellifera</Emphasis> subsp. <Emphasis Type="Italic">capensis</Emphasis> Esch.) has varied effects on nectar-feeding bird communities in South African fynbos vegetation

Outside their natural range, honeybees (Apis mellifera) are known to have detrimental effects on indigenous pollinators through exploitative or interference competition, but little is known about the effect of honeybee farming in areas where honeybees occur naturally. In the Cape Floristic Region of South Africa, where honeybees are indigenous, managed hives potentially elevate the abundance of honeybees far above natural levels, but impacts on other floral resource-dependent species have not been studied. Here we use experimental manipulation of honeybee density to test whether honeybee farming affects nectar-feeding birds. We selected the common sugarbush (Protea repens), utilized by both birds and bees, and analysed the time (before/after) by treatment (control/experiment) interaction to explore changes in bee abundance, nectar availability and bird abundance at three sites. Hive addition increased honeybee abundance in inflorescences of P. repens above expected levels. Despite experimental increase in honeybee numbers, there is no reduction in nectar sugar availability relative to the control areas. Where honeybee density was highest, sugarbird (Promerops cafer) numbers declined relative to expected, but sunbirds (Nectarinidae) were not affected at any of the sites. We conclude that stocking rates of more than one honey bee per P. repens inflorescence have detrimental effects on bird abundance due to interference, rather than resource competition.     

The effect of essential oils of sweet fennel and pignut on mortality and learning in africanized honeybees (Apis mellifera L.) (Hymenoptera: Apidae)

It was recently discovered that exposure to small concentrations of the essential oils of sweet fennel (Foeniculum vulgare Mill) or pignut [Hyptis suaveolens (L.) Poit] can be used to control aphids. What is not known is whether these oils also influence honeybee behavior. Experiments using both harnessed and free-flying foragers at concentrations used to control aphids showed that bees readily associated the odors with a reward, discriminated between them, and were not repelled. Honeybees, however, would not consume the oils when mixed with sucrose to create an unconditioned stimulus. An experiment in which harnessed bees consumed various concentrations showed that concentrations greater than 50% were detrimental. The experiments reported here provide further evidence supporting the use of conditioning techniques to evaluate the use of essential oils on honey bee behavior.     

基于TaqMan探针的蜜蜂囊状幼虫病病毒荧光PCR检测方法的建立和应用

为建立快速有效的蜜蜂囊状幼虫病检疫方法, 依据TaqMan荧光标记探针技术原理, 针对蜜蜂囊状幼虫病病毒保守序列, 设计出一对特异性引物和一条探针, 建立了一种快速检测蜜蜂囊状幼虫病病毒的荧光PCR方法。该方法对蜜蜂囊状幼虫病的检测具有较好的特异性, 与蜜蜂急性麻痹病病毒、蜜蜂慢性麻痹病病毒、蜜蜂残翼病病毒和黑蜂王台病病毒之间均无交叉反应。检测灵敏度可达1.0×10²拷贝/μL阳性质粒, 可对低病毒含量的样品进行准确检测。重复性和稳定性试验结果显示, 变异系数为1.6%, 说明该方法具有较好的重复性和稳定性。应用该方法对蜜蜂及蜂制品进行检测, 结果显示所建立的荧光PCR检测方法4 h内即可报告检测结果, 该方法具有快速、灵敏、特异及重复性好等优点, 适用于蜜蜂及其制品中蜜蜂囊状幼虫病病毒的快速检疫。     

THE KINETICS OF DARK ADAPTATION

1. Data are presented for the dark adaptation of four species of animals. They show that during dark adaptation the reaction time of an animal to light of constant intensity decreases at first rapidly, then slowly, until it reaches a constant minimum. 2. On the assumption that at all stages of adaptation a given response to light involves a constant photochemical effect, it is possible to describe the progress of dark adaptation by the equation of a bimolecular reaction. This supposes, therefore, that dark adaptation represents the accumulation within the sense cells of a photosensitive material formed by the chemical combination of two other substances. 3. The chemical nature of the process is further borne out by the fact that the speed of dark adaptation is affected by the temperature. The velocity constant of the bimolecular process describing dark adaptation bears in Mya a relation to the temperature such that the Arrhenius equation expresses it with considerable exactness when µ = 17,400. 4. A chemical mechanism is suggested which can account not only for the data of dark adaptation here presented, but for many other properties of the photosensory process which have already been investigated in these animals. This assumes the existence of a coupled photochemical reaction of which the secondary, "dark" reaction is catalyzed by the products of the primary photochemical reaction proper. This primary photochemical reaction itself is reversible in that its main products combine to form again the photosensitive material, whose concentration controls the behavior of the system during dark adaptation.     

Neural adaptation to tone sequences in the songbird forebrain: patterns, determinants, and relation to the build-up of auditory streaming

Neural responses to tones in the mammalian primary auditory cortex (A1) exhibit adaptation over the course of several seconds. Important questions remain about the taxonomic distribution of multi-second adaptation and its possible roles in hearing. It has been hypothesized that neural adaptation could explain the gradual “build-up” of auditory stream segregation. We investigated the influence of several stimulus-related factors on neural adaptation in the avian homologue of mammalian A1 (field L2) in starlings (Sturnus vulgaris). We presented awake birds with sequences of repeated triplets of two interleaved tones (ABA–ABA–…) in which we varied the frequency separation between the A and B tones (ΔF), the stimulus onset asynchrony (time from tone onset to onset within a triplet), and tone duration. We found that stimulus onset asynchrony generally had larger effects on adaptation compared with ΔF and tone duration over the parameter range tested. Using a simple model, we show how time-dependent changes in neural responses can be transformed into neurometric functions that make testable predictions about the dependence of the build-up of stream segregation on various spectral and temporal stimulus properties.     

Transgenerational phenotypic plasticity of diapause induction and related fitness cost in a commercial strain of the parasitoid Aphidius ervi Haliday

Diapause is an adaptation that insects have evolved to synchronize their life cycle with that of seasonal climatic changes and resources availability. However, cues for its induction are not always clear and, in some cases, a maternal effect may be involved. At the population level, just a part of the individuals may exhibit diapause with important consequences in terms of winter survival. Moreover, clear indicators of diapause state are difficult to identify. Diapause induction was thus investigated in the aphid parasitoid species Aphidius ervi Haliday (Hymenoptera: Braconidae) developing in the aphid Sitobion avenae (Hemiptera: Aphididae) at four crossed photothermal regimes (16 °C and 8 °C, 16:8 h L:D and 8:16 h L:D), and during 2 successive generations. We analyzed the reliability of changes in mummy color to assess for the diapausing state compared to dissections, and we measured parasitoid morphological and physiological traits. We observed that the proportion of dark brown mummies increased after one generation under low photothermal regime compared to other regimes. No diapause was recorded at 16 °C, 16:8 h L:D, while we observed 16.2% and 67.5% diapause incidence at 8 °C, 8:16 h L:D, at 1st and 2nd generation, respectively. Diapause induction is thus increased by short day-length conditions and low temperatures as well as by maternal effects. All parasitoid life-history traits (weight, size, fat content, water content, egg-load, and longevity) were affected by the photothermal regime and/or the generation. These results raise new questions on the environmental thresholds needed to induce diapause and on survival and adaptation potential of commercially available parasitoid strains in different environments.