Das Landschaftsgedächtnis der Bienen. Kleine Gehirne – überraschendes Orientierungsvermögen

We have used a new device, the harmonic radar, to monitor continously the flight paths of bees. Bees are well oriented within the area they have explored during their orientation flights. Bees transported and released at an unexpected site within this area are not lost but fly back to the hive on direct routes after a short search phase. Bees that have been trained to a feeding place may fly first to the feeding place and then back to the hive indicating that they are able to decide between two destinations for their fast return flights. Since bees could not use a beacon at the indicated goals or the structure of the horizon we conclude that their navigation memory is organized according to a geometric map.     

Orientation flight behavior in honeybees released from open or covered containers — after or without displacement

Bees captured at the feeding site were released either at a novel site or at the site at which they had been captured. Containers used during transport were either open or covered, thus allowing or preventing, respectively, free view of the panorama. Shortly before release, each bee, regardless of the covering mode used during displacement, was either (i) granted a view of the release site, (ii) allowed to see only overall brightness by covering the container with translucent drawing paper, or (iii) denied any view of the release site. Orientation flights performed by each bee upon release were recorded. Irrespective of whether they had been transported in open or covered containers, bees granted a view of the release site prior to release usually flew away without performing orientation flights, whereas bees prevented from viewing the release site performed intensive orientation flights. An intermediate intensity of orientation flights was displayed by bees leaving translucent containers. Assuming that the intensity of orientation flight is a measure of the state of orientation of the released bee, we conclude that view of the release site, but not view of the environment during displacement, is crucial for orientation.Dedicated to Wolfgang Wickler on occasion of the 65th anniversary of his birthday.     

Role of moth ocelli in timing flight initiation at dusk

ABSTRACT. The role of ocelli in timing flight initiation of cabbage looper moths, Trichoplusia ni (Hübner) (Noctuidae), at dusk was studied under simulated sunset conditions using a multichannel actograph. The mean time of flight initiation was determined for control, sham (sham/occluded, sham ablated, unilateral ablated) and experimental (ocellus occluded, ocellus ablated moths). Ocellus occlusion delayed flight initiation on the first day following treatment but was less effective on the subsequent days. Ocellus ablation also delayed flight initiation on the first day, and also produced pronounced delays on subsequent days. In studies where the sunset was advanced 1 h, control sham ocellus occluded and unilaterally ocellus ablated moths responded to the advance, but bilaterally ocellus ablated moths did not. These results indicate that moths make use of input from the ocelli in determining the threshold light intensity for flight and in making adjustments to small light-phase changes.     

Visual gaze control during peering flight manoeuvres in honeybees

As animals travel through the environment, powerful reflexes help stabilize their gaze by actively maintaining head and eyes in a level orientation. Gaze stabilization reduces motion blur and prevents image rotations. It also assists in depth perception based on translational optic flow. Here we describe side-to-side flight manoeuvres in honeybees and investigate how the bees’ gaze is stabilized against rotations during these movements. We used high-speed video equipment to record flight paths and head movements in honeybees visiting a feeder. We show that during their approach, bees generate lateral movements with a median amplitude of about 20 mm. These movements occur with a frequency of up to 7 Hz and are generated by periodic roll movements of the thorax with amplitudes of up to ±60°. During such thorax roll oscillations, the head is held close to horizontal, thereby minimizing rotational optic flow. By having bees fly through an oscillating, patterned drum, we show that head stabilization is based mainly on visual motion cues. Bees exposed to a continuously rotating drum, however, hold their head fixed at an oblique angle. This result shows that although gaze stabilization is driven by visual motion cues, it is limited by other mechanisms, such as the dorsal light response or gravity reception.     

Thorax vibrations of a stingless bee (<Emphasis Type="Italic">Melipona seminigra</Emphasis>). I. No influence of visual flow

An important question in stingless bee communication is whether the thorax vibrations produced by foragers of the genus Melipona upon their return to the nest contain spatial information about food sources or not. As previously shown M. seminigra is able to use visual flow to estimate flight distances. The present study investigated whether foraging bees encode the visually measured distance in their thorax vibrations. Bees were trained to collect food in flight tunnels lined with a black-and-white pattern on their side walls and floor, which substantially influenced the image motion they experienced. When the bees had collected inside the tunnels the temporal pattern of their vibrations differed significantly from the pattern after collecting in a natural environment. These changes, however, were not associated with the visual flow experienced inside the tunnel. Bees collecting in tunnels offering little visual flow (stripes parallel to flight direction) modified their vibrations similarly to bees collecting in tunnels with high image motion (cross stripes). A higher energy expenditure due to drastically reduced flight velocities inside the tunnel is suggested to be responsible for changes in the thorax vibrations. The bees' vibrations would thus reflect the overall energetic budget of a foraging trip.     

The role of the ocelli in the phototactic behaviour of the haematophagous bug Triatoma infestans

In addition to compound eyes, most adult insects posses two or three simple eyes, the ocelli. The function of these photoreceptors remains elusive in most cases. Triatomine bugs posses two well-developed ocelli, located in a latero-dorsal position, behind the compound eyes. We tested the role of the ocelli in the phototactic behaviour of Triatoma infestans, by measuring the time spent by adult males in the dark half of an experimental arena, which had the other half illuminated. The occlusion of the ocelli or the compound eyes alone had little effect on the phototactic response of the bugs. Only those insects which had both their ocelli and compound eyes occluded showed a significant reduction in their negative response to light. The ability of the ocelli of T. infestans to mediate the phototactic response by themselves (i.e., not through the modulation of compound eyes sensitivity) constitutes the first report on this function in insects.     

Hornets Can Fly at Night without Obvious Adaptations of Eyes and Ocelli

Hornets, the largest social wasps, have a reputation of being facultatively nocturnal. Here we confirm flight activity of hornet workers in dim twilight. We studied the eyes and ocelli of European hornets (Vespa crabro) and common wasps (Vespula vulgaris) with the goal to find the optical and anatomical adaptations that enable them to fly in dim light. Adaptations described for obligately nocturnal hymenoptera such as the bees Xylocopa tranquebarica and Megalopta genalis and the wasp Apoica pallens include large ocelli and compound eyes with wide rhabdoms and large facet lenses. Interestingly, we did not find any such adaptations in hornet eyes or ocelli. On the contrary, their eyes are even less sensitive than those of the obligately diurnal common wasps. Therefore we conclude that hornets, like several facultatively nocturnal bee species such as Apis mellifera adansonii, A. dorsata and X. tenuiscapa are capable of seeing in dim light simply due to the large body and thus eye size. We propose that neural pooling strategies and behavioural adaptations precede anatomical adaptations in the eyes and ocelli when insects with apposition compound eyes turn to dim light activity.     

凹唇壁蜂成蜂体重与取食对其飞行能力的影响

凹唇壁蜂Osmia excavata Alfken被广泛应用于我国北方果树的传粉, 而其飞行能力是影响其传粉效率的重要因素。本研究通过飞行磨吊飞试验, 评估了凹唇壁蜂雌蜂和雄蜂飞行能力的差异以及取食对其飞行能力的影响。结果表明, 凹唇壁蜂雌蜂体重（116.30 mg）显著大于雄蜂（59.80 mg） (P<0.001), 雌蜂的最大飞行速度（3.44 km/h）显著大于雄蜂（2.36 km/h）, 雄蜂的飞行距离和最大飞行速度与其飞行前体重成显著的正相关性, 雌蜂的飞行时间与其飞行前体重成显著正相关性, 而雌蜂的平均飞行速度与其体重成显著负相关性; 雌蜂的日平均飞行距离为0.23 km, 根据雌蜂以巢为中心, 采集花粉繁殖后代的生物学习性, 蜂巢之间的放置距离应少于100 m。取食蜂蜜后, 雌雄壁蜂的飞行距离、 飞行时间、 最大飞行速度均有提高的趋势, 建议在田间应用时, 可在蜂巢附近放置蜂蜜或种植其他蜜源植物给初羽化的凹唇壁蜂提供食物补充能量。本研究明确了雌、 雄壁蜂的飞行能力和出茧后补充食物对于壁蜂飞行的促进作用, 为有效地利用凹唇壁蜂进行传粉提供了理论依据。     

Simulations of a model for transmitter kinetics

The influence of the internal water balance on the phototactic behaviour in the walking female fly (Calliphora erythrocephala Meig.) was investigated. The phototactic reaction depends on the age of the flies and the duration of water withdrawal. In young blowflies with progressive dehydration, the strength of the light reaction varies considerably from fly to fly. From the 4th. day of life onwards up to day 21 the flies respond much more homogeneously and elicit a reproduceable temporal pattern of reaction (Figs. 2 and 3). All the following statements refer to the behaviour of 10-day-old, virgin females, which, under optimal humidity conditions, have been shown to be spontancously photonegative (Meyer, 1978). The phototactic reaction of progressively dehydrated flies depends in a characteristic manner on the illumination conditions during the intervals between tests. If the flies are kept in darkness during these intervals, the light reaction varies rhythmically, with a period of almost exactly 12 h (Figs. 4a and 5). Under the test conditions this rhythm is found not to vary with the time of day (Fig.4a), or with the length of the between-test intervals, for intervals up to 4h long (Fig. 6). If the flies are kept under illumination during the intervals between tests, the light reaction becomes arhythmical. After an initial maximum after 2–4h of dehydration, further photopositive responses are severely suppressed (Fig. 4b). When the ocelli are covered, the between-test illumination no longer influences the mean response to light. The arhythmic dehydrationtime vs. light-reaction curve in this case is characterised by a strong sustained enhancement of runs towards the light after 10h of dehydration (Fig. 7). A preliminary model of a possible control system for this moisture-dependent phototactic switching is presented, from which all essential results can be deduced. This system determines the phototactic turning direction from the ocelli afferences. These afferences act upon the central nervous system in two ways: directly and also indirectly via the internal water regulation.This work was supported by the DFG-(Me417/4)     

Bee vision of pattern and 3D. The Bidder Lecture 1994

Insect vision is nothing if not active. The regular head movements, called saccades, enable the fly Drosophila to keep a straight path in flight despite inequalities in the thrust of the wings. Using their own motion, bees in flight measure the ranges of nearby objects. A long history of research shows that bees discriminate visually in ways that depend on their activity or task, so we must distinguish between vision during flying, fixating or hovering and landing. Bees return again and again for a reward of sugar solution and use their eyes to find their way. In an apparatus that makes them discriminate between two simulataneously visible but regularly interchanged targets, seen at a distance of 27 cm, bees are able to distinguish a remarkable number of simple patterns, but they fail in certain critical cases. The results can be explained with the hypothesis that bees have several broadly tuned overlapping filters with large fields that respond to the predominant orientation in a region of the image, and others for radial and circular patterns. Together with colour, these filters are independent of range. Bees prefer to use landmarks where they can, then global pattern at the largest scale, and lastly the detail around the goal. The way that discrimination of one visual feature is independent of other variables can be explained by models analogous to the colour triangle in colour discrimination.     

Tetragonisca guard bees interpret expanding and contracting patterns as unintended displacement in space

Guard bees of the stingless bee Tetragonisca angustula (Apidae: Meliponinae) hover in stable positions in front of the nest to protect the flight corridor leading to the nest entrance against insect intruders. To unravel the visual control of station keeping, we exposed these hovering guards to expanding and contracting patterns at the nest front. The bees fly away from an expanding pattern and towards the centre of a contracting pattern along a line connecting their initial position and the centre of expansion regardless of where in the visual field they view the pattern. The response of bees to a spinning radial pattern is different: they fly parallel to the pattern, up and down or forward and backward depending on whether they initially hover to the side, above or below the centre of rotation. The bees respond to horizontal and to vertical expansion and contraction. They also adjust their distance relative to a rotating spiral which produces a realistic flow field and thus allowed us to test to what extent the bees minimize image motion speed. We find that guard bees indeed move in the appropriate direction to minimize the image motion speed they experience. A comparison of bees hovering at different distances from the nestfront at the onset of pattern motion and experiencing very different image velocities shows that the dynamics of the reaction is quite uniform. At the pattern velocities tested, we did not find evidence that guard bees use image motion to control their flight speed. The bees' response rather suggests that the underlying mechanism might be insensitive to the size of motion vectors. Accepted: 2 April 1997     

Visual ecology of Indian carpenter bees I: Light intensities and flight activity

Bees are mostly active during the daytime, but nocturnality has been reported in some bee families. We studied temporal flight activity in three species of carpenter bees (genus Xylocopa) in relation to light intensities. X. leucothorax is diurnal, X. tenuiscapa is largely diurnal being only occasionally crepuscular, while X. tranquebarica is truly nocturnal. Occasional forays into dim light by X. tenuiscapa are likely to be due to the availability of richly rewarding Heterophragma quadriloculare (Bignoniaceae) flowers, which open at night. X. tranquebarica can fly even during the moonless parts of nights when light intensities were lower than 10⁻⁵ cd m⁻², which makes this species the only truly nocturnal bee known so far. Other known dim-light species fly during crepuscular or moonlit periods. We compare eye and body sizes with other known diurnal and dim-light bees. We conclude that while extremely large ocellar diameters, large eye size:body size ratio, large number of ommatidia and large ommatidial diameters are all adaptations to dim-light foraging, these alone do not sufficiently explain the flights of X. tranquebarica in extremely dim light. We hypothesise that additional adaptations must confer extreme nocturnality in X. tranquebarica.     

A stingless bee (Melipona seminigra) uses optic flow to estimate flight distances

Foragers of a stingless bee, Melipona seminigra, are able to use the optic flow experienced en route to estimate flight distance. After training the bees to collect food inside a flight tunnel with black-and-white stripes covering the side walls and the floor, their search behavior was observed in tunnels lacking a reward. Like honeybees, the bees accurately estimated the distance to the previously offered food source as seen from the sections of the tunnel where they turned around in search of the food. Changing the visual flow by decreasing the width of the flight tunnel resulted in the underestimation of the distance flown. The removal of image motion cues either in the ventral or lateral field of view reduced the bees' ability to gauge distances. When the feeder inside the tunnel was displaced together with the bees feeding on it while preventing the bee from seeing any image motion during the displacement the bees experienced different distances on their way to the food source and during their return to the nest. In the subsequent test the bees searched for the food predominantly at the distance associated with their return flight.     

Honey bee orientation behaviour and the influence of flower distribution on foraging movements

ABSTRACT.

• 1 Directional movement by foraging honey bees (Apis mellifera L.) was studied on several flower arrays. The most frequent move among equidistant flower stalks was straight ahead from stalk to stalk with frequencies decreasing for increasing turn angles. Turns to the left were about equal in frequency to turns to the right.
• 2 Bees maintained directionality when moving from flower stalks that had been rotated 90° counterclockwise while the bee was on the stalk (no difference between moves from rotated stalks and unrotated controls). Thus, directionality is maintained by the bee and is not an artefact of flower distribution.
• 3 Bees also maintained directionality when the entire array was rotated around the flower stalk the bee was on. Thus, bees use an external cue to orientate in a given direction rather than fixing on an inflorescence within the flower array.
• 4 Bees foraging on very different flower arrays differed in patterns of directionality and in distances flown between flower stalks. Therefore, even though bees maintain directionality using external cues, flower distribution can nevertheless influence flight patterns.

     

Nocturnal orientation by the Asian honey bee, Apis dorsata

Honey bees have been observed to forage and dance on moonlit nights, but it has never been established whether the moon serves as a reference in orienting nocturnally active bees. The present study, of the Asian honey bee Apis dorsata, suggests that although the moon's illumination is essential for nocturnal flight, the moon itself is ignored for orienting the dances. Rather, bees probably use the sun's position as a reference point for their dances, even though the sun is below the horizon. This ability may involve an extension of the mechanism that honey bees employ to find the sun on overcast days.     

Bees exposed to climate change are more sensitive to pesticides

Bee populations are exposed to multiple stressors, including land-use change, biological invasions, climate change, and pesticide exposure, that may interact synergistically. We analyze the combined effects of climate warming and sublethal insecticide exposure in the solitary bee Osmia cornuta. Previous Osmia studies show that warm wintering temperatures cause body weight loss, lipid consumption, and fat body depletion. Because the fat body plays a key role in xenobiotic detoxification, we expected that bees exposed to climate warming scenarios would be more sensitive to pesticides. We exposed O. cornuta females to three wintering treatments: current scenario (2007–2012 temperatures), near-future (2021–2050 projected temperatures), and distant-future (2051–2080). Upon emergence in spring, bees were orally exposed to three sublethal doses of an insecticide (Closer, a.i. sulfoxaflor; 0, 4.55 and 11.64 ng a.i./bee). We measured the combined effects of wintering and insecticide exposure on phototactic response, syrup consumption, and longevity. Wintering treatment by itself did not affect winter mortality, but body weight loss increased with increasing wintering temperatures. Similarly, wintering treatment by itself hardly influenced phototactic response or syrup consumption. However, bees wintered at the warmest temperatures had shorter longevity, a strong fecundity predictor in Osmia. Insecticide exposure, especially at the high dose, impaired the ability of bees to respond to light, and resulted in reduced syrup consumption and longevity. The combination of the warmest winter and the high insecticide dose resulted in a 70% longevity decrease. Smaller bees, resulting from smaller pollen–nectar provisions, had shorter longevity suggesting nutritional stress may further compromise fecundity in O. cornuta. Our results show a synergistic interaction between two major drivers of bee declines, and indicate that bees will become more sensitive to pesticides under the current global warming scenario. Our findings have important implications for pesticide regulation and underscore the need to consider multiple stressors to understand bee declines.     

Ocellar optics in nocturnal and diurnal bees and wasps

Nocturnal bees, wasps and ants have considerably larger ocelli than their diurnal relatives, suggesting an active role in vision at night. In a first step to understanding what this role might be, the morphology and physiological optics of ocelli were investigated in three tropical rainforest species – the nocturnal sweat bee Megalopta genalis, the nocturnal paper wasp Apoica pallens and the diurnal paper wasp Polistes occidentalis – using hanging-drop techniques and standard histological methods. Ocellar image quality, in addition to lens focal length and back focal distance, was determined in all three species. During flight, the ocellar receptive fields of both nocturnal species are centred very dorsally, possibly in order to maximise sensitivity to the narrow dorsal field of light that enters through gaps in the rainforest canopy. Since all ocelli investigated had a slightly oval shape, images were found to be astigmatic: images formed by the major axis of the ocellus were located further from the proximal surface of the lens than images formed by the minor axis. Despite being astigmatic, images formed at either focal plane were reasonably sharp in all ocelli investigated. When compared to the position of the retina below the lens, measurements of back focal distance reveal that the ocelli of Megalopta are highly underfocused and unable to resolve spatial detail. This together with their very large and tightly packed rhabdoms suggests a role in making sensitive measurements of ambient light intensity. In contrast, the ocelli of the two wasps form images near the proximal boundary of the retina, suggesting the potential for modest resolving power. In light of these results, possible roles for ocelli in nocturnal bees and wasps are discussed, including the hypothesis that they might be involved in nocturnal homing and navigation, using two main cues: the spatial pattern of bright patches of daylight visible through the rainforest canopy, and compass information obtained from polarised skylight (from the setting sun or the moon) that penetrates these patches.     

The Grooming Invitation Dance of the Honey Bee

The grooming invitation dance is a striking behavior in honey bee colonies that has not been extensively studied. The objectives of this study were (1) to describe the dance through video analysis, (2) to test the functional hypothesis that it is a grooming solicitation signal, and (3) to analyze the stimuli that cause its production. A worker bee producing the grooming invitation dance stands stationary and vibrates her whole body from side‐to‐side at a frequency of 4.2 ± 0.2 Hz for 9.3 ± 1.0 s. Sometimes the bee mixes bouts of body vibration with brief bouts of self‐grooming (average duration = 1.4 s). Bees that perform the grooming invitation dance have a far higher probability of being quickly groomed by a nest mate than do bees that do not perform the dance. Bees that had chalk dust puffed onto the bases of their wings produced significantly more grooming invitation dances than did control bees that received only puffs of air. This shows that it may be the accumulation of small particles at the bases of the wings that normally triggers the dance. We suggest that the evolutionary origin of this signal is self‐grooming behavior.     

A Mathematical Model for Flight Guidance in Honeybee Swarms

When a colony of honeybees relocates to a new nest site, less than 5?% of the bees (the scout bees) know the location of the new nest. Nevertheless, the small minority of informed bees manages to provide guidance to the rest and the entire swarm is able to fly to the new nest intact. The streaker bee hypothesis, one of the several theories proposed to explain the guidance mechanism in bee swarms, seems to be supported by recent experimental observations. The theory suggests that the informed bees make high-speed flights through the swarm in the direction of the new nest, hence conspicuously pointing to the desired direction of travel. This work presents a mathematical model of flight guidance in bee swarms based on the streaker bee hypothesis. Numerical experiments, parameter studies, and comparison with experimental data are presented.     

Pollen in Bee-Flower Relations Some Considerations on Melittophily*

Pollen and nectar are usually lumped together as floral rewards for pollinating bees, but they play totally different roles for flowers and bees (Table 1), as well as in the relationship between them. While flowers are specialized for certain pollinators via nectar, bees specialize on certain flowers via pollen. While flowers need pollen as a prerequisite for pollination, it is the essential larval food in bees. Thus, there is a strong competition between them for pollen. Foraging for pollen must be divided into three phases: uptake in the flower, reloading into and homeward transport within a carrying container. Bees have specializations for transport but hardly any for pollen uptake - and thus for pollination. Bees actively harvesting pollen usually do not pollinate. This only happens as a consequence of contamination of the bee by pollen. From these data a scenario is provided for the evolution of bees and bee flowers. Specialized bee flowers are often characterized by their ability to hide pollen from the bees and at the same time use them as optimal pollinators. If the relationship of bees and flowers is mutualistic at all it is best described as a balanced mutual exploitation.