Descriptive study of antennation during trophallactic unloading contacts in honeybees <Emphasis Type="Italic">Apis mellifera carnica</Emphasis>

Summary  Trophallaxis, the mouth-to-mouth transfer of food, is a widespread behavior occurring between individuals of eusocial insect societies. Antennal movements during food transfer are, in honeybees, too rapid to be characterized using standard video recordings. Using a high-speed camera (200 frames/s), we recorded nectar unloading performed by forager honeybees (Apis mellifera carnica Spinola) within the hive once they returned from collecting sugar solution at a feeder that delivered nectar at a variable rate. Frequency patterns attained a mean value of 13 Hz. Antennation intensity showed a tendency to increase with the reward rate recently exploited by the food donor. This raises the question whether or not antennation intensity is a reliable parameter providing modulatory information related to food-source profitability.Received 12 September 2002; revised 21 March 2002; accepted 24 April 2003.     

Discrimination of nestmate workers and drones in honeybees

Summary Discrimination of nestmates from non-nestmates has mainly been investigated in female social insects. Little is known about discrimination of males. Here we show that under natural conditions at the nest entrance, honeybee workers can discriminate nestmate drones from non-nestmate drones as effectively as they can discriminate nestmate workers from non-nestmate workers.     

Pattern recognition in honeybees: chromatic properties of orientation analysis

The spectral properties of the discrimination of pattern orientation in freely flying honeybees (Apis mellifera) were examined. Bees were trained to discriminate between two random black/white gratings oriented perpendicularly to each other, one of which was associated with a reward. Subsequently the bees were tested on two-colour gratings or gratings consisting of grey and coloured stripes, providing a range of different chromatic contrasts, luminance contrasts and specific channel contrasts. The results of these experiments indicate that orientation analysis in the honeybee is mediated almost exclusively by the green receptor channel, although the bee's visual system as a whole is endowed with excellent trichromatic colour vision.     

Pattern recognition in honeybees: eidetic imagery and orientation discrimination

The roles of eidetic imagery and orientational cues, respectively, in the discrimination of visual patterns by honeybees (Apis mellifera) were evaluated by training the bees to discriminate between patterns consisting of periodic, black and white square wave gratings. Training and tests with a number of different pairs of patterns revealed that bees use orientational cues almost exclusively, if such are present, and make use of eidetic images only when orientational cues are not available. On the other hand, if a pattern carries strong orientational cues, bees learn the orientation even if it is irrelevant to the discrimination task on which they are trained.     

Detection of coloured stimuli by honeybees: minimum visual angles and receptor specific contrasts

Honeybees Apis mellifera were trained to distinguish between the presence and the absence of a rewarded coloured spot, presented on a vertical, achromatic plane in a Y-maze. They were subsequently tested with different subtended visual angles of that spot, generated by different disk diameters and different distances from the decision point in the device. Bees were trained easily to detect bee-chromatic colours, but not an achromatic one. Chromatic contrast was not the only parameter allowing learning and, therefore, detection: _min, the subtended visual angle at which the bees detect a given stimulus with a probability P ₀ = 0.6, was 5° for stimuli presenting both chromatic contrast and contrast for the green photoreceptors [i.e. excitation difference in the green photoreceptors, between target and background (green contrast)], and 15° for stimuli presenting chromatic but no green contrast. Our results suggest that green contrast can be utilized for target detection if target recognition has been established by means of the colour vision system. The green-contrast signal would be used as a far-distance signal for flower detection. This signal would always be detected before chromatic contrast during an approach flight and would be learned in compound with chromatic contrast, in a facilitation-like process.     

Genetic variance of mating frequency in the honeybee (Apis mellifera L.)

Summary. The genetic variance of queen mating frequency was studied in honeybees (Apis mellifera carnica). Worker offspring (N = 966) of 28 naturally mated half sister-queens (r = 0.25) from seven unrelated breeding lines were genotyped at four DNA microsatellites. The mating frequencies of the queens were derived from the offspring genotypes. The number of observed matings per queen ranged from 10 to 28 with an average of 17.32 ± 1.10 (number of estimated matings: 24.94 ± 2.51; number of effective matings: 20.09 ± 1.73). Half-sib analyses of the breeding lines were used to estimate heritability. Heritability was h² = 0.449 ± 0.135 for the estimated number of matings and h² = 0.262 ± 0.103 for the number of effective males, which are both significantly different from zero. We conclude that a high genetic variance for polyandry in honeybees can be favored by balanced selection between individual queen and colony level.Received 16 October 2003; revised 4 May 2004; accepted 4 May 2004.     

Hearing in honeybees: localization of the auditory sense organ

Airborne sound signals emitted by dancing honeybees (Apis mellifera) contain information about the locations of food sources. Honeybees can perceive these near field sounds and rely on them to decode the messages of the dance language. The dance sound is characterized by rhythmical air particle movement of high velocity amplitudes. The aim of the present study was to identify the sensory structures used to detect near field sound signals. In an operant conditioning experiment, bees were trained to respond to sound. Ablation experiments with these trained bees revealed that neither mechanosensory hairs on the antennae or head nor bristle fields at the joints of the antenna, but Johnston's organ, a chordotonal organ in the pedicel of the antenna, is used to detect near field sound in honeybees.     

Thermal behavior of round and wagtail dancing honeybees

The thermal behavior of round and wagtail dancing honeybees (Apis mellifera carnica) gathering sucrose solutions of concentrations between 0.5 and 2 mol·l^-1 was investigated under field conditions by infrared thermography (30–506 m flight distance). During the stay inside the hive thoracic surface temperature ranged from 31.4 to 43.9 °C. In both round and wagtail dancing honeybees the concentration of sucrose in the food influenced dancing temperature in a non-linear way. Average dancing temperature was 37.9 °C in foragers gathering a 0.5 mol·l^-1 sucrose solution, 40.1°C with a 1 mol·l^-1, 40.6°C with a 1.5 mol·l^-1 and 40.7°C with a 2 mol·l^-1 solution. The variability of thoracic temperature was highest with the 0.5 mol·l^-1 and lowest with the 1.5 and 2 mol·l^-1 concentrations. Thoracic temperatures during trophallactic contact with hive bees were similar to dancing temperature at 1.5 mol·l^-1 but lower at the other concentrations. During periods of distribution of food to hive bees (trophallactic contact >2.5s) the dancers' thorax cooled down by more than 0.5°C considerably more frequently with the 0.5 mol·l^-1 solution (65% of cases) than with the 1.5 mol·l^-1 solution (26%). By contrast, heating the thorax up by more than 0.5°C was infrequent with the 0.5 mol·l^-1 solution (2%) but occurred at a maximum rate of 26% with the 1.5 mol·l^-1 solution. Bees gathering the 1 or 2 mol·l^-1 solutions showed intermediate behavior. Linear model analysis showed that at higher concentrations the dancers compensated better for variations of hive air temperature: per 1 °C increase of hive temperature dancing temperature increased by 0.34, 0.22, 0.12, and 0.13 °C with 0.5, 1, 1.5, and 2 mol·l^-1 sucrose solutions, respectively. The results furnish evidence that dancing honeybees follow a strategy of selective heterothermy by tuning their thermal behavior to the needs of the behavior performed at the moment. Thoracic temperature is regulated to a high level and more accurately when fast exploitation of profitable food sources is recommended. Thoracic temperature is lowered when the ratio of gain to costs of foraging becomes more unfavorable.Abbreviations SD standard deviation - SD _reg SD around regression line - H _rel relative humidity at feeding station - T _a air temperature at feeding station - T _i air temperature near the dancers - T _d Thoracic surface temperatures - T _d dancing - T _tr trophallactic contact (distribution of food) - T _w walking - T _stay mean temperature of total stay in the hive     

Long-term effects of soybean protease inhibitors on digestive enzymes, survival and learning abilities of honeybees

We have investigated the effects of long-term ingestion of two serine proteinase inhibitors (PIs), the Kunitz Soybean trypsin inhibitor (SBTI) and the Bowman-Birk inhibitor (BBI) on survival, learning abilities involved in the foraging behaviour, and digestive physiology of the honeybee (Apis mellifera L., Hymenoptera). A threshold-dose was established, above which adverse effects of long-term ingestion of the PIs tested are to be expected. The experiments reported herein could be extended to other PIs or gene products used to confer insect resistance, and be part of a general procedure used to assess the innocuousness of transgenic melliferous plants to honeybees.     

Hearing in honeybees: operant conditioning and spontaneous reactions to airborne sound

Summary Airborne sound signals emitted by dancing bees (Apis mellifera) play an essential role in the bees' dance communication. It has been shown earlier that bees can learn to respond to airborne sounds in an aversive conditioning paradigm. Here we present a new training paradigm. A Y-choice situation was used to determine the frequency range and amplitude thresholds of hearing in bees. In addition, spontaneous reactions of bees to airborne sound were observed and used to determine thresholds of hearing. Both methods revealed that bees are able to detect sound frequencies up to about 500 Hz. The hearing threshold is 100–300 mm/s peak-to-peak velocity and is roughly constant over the range of detectable frequencies. The amplitude of the signals emitted in the dance language is 5 to 10 times higher, so we can conclude that bees can easily detect the dance sounds.     

Nosema ceranae, a new parasite in Thai honeybees

Adult workers of Apis cerana, Apis florea and Apis mellifera from colonies heavily infected with Nosema ceranae were selected for molecular analyses of the parasite. PCR-specific 16S rRNA primers were designed, cloned, sequenced and compared to GenBank entries. The sequenced products corresponded to N. ceranae. We then infected A. cerana with N. ceranae spores isolated from A. florea workers. Newly emerged bees from healthy colonies were fed 10,000, 20,000 and 40,000 spores/bee. There were significant dosage dependent differences in bee infection and survival rates. The ratio of infected cells to non-infected cells increased at 6, 10 and 14 d post infection. In addition, hypopharyngeal glands of bees from the control group had significantly higher protein concentrations than infected groups. Bees infected with 40,000 spores/bee had the lowest protein concentrations. Thus, N. ceranae isolated from A. florea is capable of infecting another bee species, impairing hypopharyngeal gland protein production and reducing bee survival in A. cerana.     

Energetics of nectar production in some strawberry cultivars as a predictor of floral choice by honeybees

Nectar productivity of thirteen cultivars of strawberry (Fragariq sp.) was examined in relation to the foraging activity ofApis cerana indica F. andApis mellifera L. The cultivars were found to vary in the quantity of nectar produced (0·020 to 0·735 ul/flower/day), nectar-sugar concentration (30 to 42%) and amount of sugar (0.006 to 0·2898 mg/flower/day). Consequently energy reward varied from 0·1037 to 4·851 joules/ flower/day. Foraging of bothApis cerana indicq andApis melliferq correlated with energy yields. The results suggest that cultivars with higher caloric rewards had a competitive edge over the others in attracting foraging populations of both species of the bee.     

The forest or the trees: preference for global over local image processing is reversed by prior experience in honeybees

Traditional models of insect vision have assumed that insects are only capable of low-level analysis of local cues and are incapable of global, holistic perception. However, recent studies on honeybee (Apis mellifera) vision have refuted this view by showing that this insect also processes complex visual information by using spatial configurations or relational rules. In the light of these findings, we asked whether bees prioritize global configurations or local cues by setting these two levels of image analysis in competition. We trained individual free-flying honeybees to discriminate hierarchical visual stimuli within a Y-maze and tested bees with novel stimuli in which local and/or global cues were manipulated. We demonstrate that even when local information is accessible, bees prefer global information, thus relying mainly on the object''s spatial configuration rather than on elemental, local information. This preference can be reversed if bees are pre-trained to discriminate isolated local cues. In this case, bees prefer the hierarchical stimuli with the local elements previously primed even if they build an incorrect global configuration. Pre-training with local cues induces a generic attentional bias towards any local elements as local information is prioritized in the test, even if the local cues used in the test are different from the pre-trained ones. Our results thus underline the plasticity of visual processing in insects and provide new insights for the comparative analysis of visual recognition in humans and animals.     

Hearing in honeybees: the mechanical response of the bee's antenna to near field sound

In the dance language, honeybees use airborne near field sound signals to inform their nestmates of the location of food sources. In behavioral experiments it has recently been shown that Johnston's organ, a chordotonal organ located in the pedicel of the antenna, is used to perceive these sound signals. In the present study the mechanical response of the antennal flagellum to stimulation with near field sound signals was investigated using laser vibrometry. The absolute amplitudes of antennal deflection with acoustical stimulation, the response to sounds of different displacement and velocity amplitudes, the shape of movement of the flagellum, the mechanical frequency response and the mechanical directional sensitivity of the auditory sense organ of the honeybee are described. Using pulsed stimuli simulating the dance sounds it is shown that the temporal pattern of the dance sound is resolved on the level of antennal vibrations.