What attracts honeybees to a waggle dancer?

In the vicinity of a dancer, a honeybee can become a dance follower after touching the dancer or a dance follower with an antenna. If the attraction occurs without such antennal contact, the strength of the attraction over distance depends on several factors: the kind of dance floor (empty open cells versus capped brood cells); whether dancers and dance followers stand on the same substratum or on separate substrata; the position and direction of the attracted bee relative to the dancer bee; the size of the dance group (the dancer plus follower bees); and the light conditions under which the dance takes place. Dances on open cells are significantly more attractive than dances on sealed cells. Dancers on open cells attracted 90% of all followers from within 27 mm (about five to six cell diameters). Dancers on sealed cells attracted 90% of all followers within 18 mm (about three cell diameters). The majority of bees that were attracted by the dancer were standing laterally to the dancer. Dances illuminated by artificial visible light are significantly more attractive than dances illuminated by infrared light. As a group, “glassplate bees” (bees standing mechanically isolated from the dancer bee) were least attracted. Accepted: 11 August 1998     

E-Vector orientation in bees

Summary If an insect is able to determine the direction of polarization in any point of the sky, this ability does not in itself guarantee that the insect can orientate unambiguously. Such would only be the case, if every point in the sky had its own exclusive direction of polarization. In thee-vector pattern of the sky, however, each direction of polarization is found at many different points. For compass orientation the insect has therefore to use some information on the geometry of thee-vector pattern in the sky. In general, eache-vector occurs twice at a given elevation (Fig. 1). The angular separation between the positions of identicale-vectors depends on the elevations of thee-vectors above the horizon and on the height of the sun. Except at sunrise and sunset, 180°.If a bee is trained to fly in a certain direction to a food source, the direction of its waggle dance on a horizontal comb points directly towards the goal (provided that the bee is able to view the sky). However, if the bee is only allowed to view a singlee-vector in the sky (or a single artificially adjustede-vector), it should perform ambiguous orientation. One expects the bee to prefer two dance directions separated by the proper angular distance . One of these two dance directions should point at the food source.The bees indeed dance in two directions. However, there are two unexpected results: (1) The angular distance between the two preferred directions invariably amounts to =180°. (2) One of the preferred directions points closely, but not exactly at the goal. What one can deduce from these single-e-vector tests is that the bee uses a rather generalized internal representation of thee-vector pattern in the sky. This paper describes the generale-vector characteristic applied by a dancing bee that only views a singlee-vector in the sky (diameter of the celestial patch or the artificially polarized light source 10°). This generale-vector characteristic of the bee (Fig. 9) more closely fits the meane-vector distribution near the zenith thane-vector distributions in other parts of the sky (Fig. 11).This article is dedicated to Prof. Dr. H. Autrum in honor of his seventieth birthdayThe research has been supported by Swiss National Science Foundation Grant 3.814.72, continued by Grant 3.529.75, and by the Academy of Science and Literature at Mainz. We would like to thank Dr. R. Schinz (Purdue University) for cooperation and fruitful discussions as well as Mrs. V. Güttinger, Mrs. A. Rossel-Jäckle, and Miss A. Blischke for technical assistance.     

Honey bees: Photoreceptors participating in orientation behaviour to light and gravity

Summary The ability of honey bees to dance compromise directions when confronted with conflicting light and gravity references was used to determine their spectral sensitivity. The action spectrum has peaks at 450 nm and 550 nm and thus indicates the contributions of the blue and green receptors of the bees' compound eyes. There is no obvious contribution from the UV receptors. The dance directions indicate that blue and yellow-green light is regarded as sunlight. UV light seems not to be so interpreted, though it clearly influences the bees' orientation to gravity, if it is polarized. This result is consistent with our earlier findings which demonstrated that the bees' 350 nm receptors are used as detectors of blue skylight rather than sunlight. Because the receptor contributions differ, there is a clear distinction between the sun compass behaviour apparent in the dances and the phototaxis responses observed in other contexts.Supported by the Deutsche Forschungsgemeinschaft, DFG-Schwerpunkt 322, 243     

Processing of sun-azimuth information by honey bees

The dance language of honey bees (Apis mellifera) indicates with good accuracy the distance and direction of a food source. When the direction of a food source is moved, however, the dances of returning foragers indicate the former location rather than the new direction just visited. After subsequent trips the dancers slowly accommodate to the change, and full compensation is achieved only after about 40 min. These and other equally curious results suggest that a running-average processing system may be at work in honey bee navigation.     

Polarized light orientation of the honey bee: The minimum visual angle

Summary A field of linearly polarized light less than one degree in diameter suffices for a honey bee to orient its dance on a horizontal comb. Thus at any one moment, a surprisingly small part of the honey bee's complex eye, consisting of 3 to 7 ommatidia, is sufficient for perception of thee-vector direction of polarized light.     

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.     

The waggle dance of the honey bee: Which bees following a dancer successfully acquire the information?

During the waggle dance of the honeybee, the dancer is able to tell her nestmates the distance and direction to a rich food source (Frisch, 1967). Little is known about how waggle dance followers are able to read the waggle dance in the darkness of a hive. Initial observations showed that not all of the bees that appear to be dance followers behave the same. Some bees maneuver themselves behind the dancer, while others do not. The paths of a single dancer, trained to an artificial food source, and her followers were traced during the waggle runs. The success of these dance followers was compared to their position relative to the dancer. The results of this study show that during a waggle run a dance follower must position itself within a 30° arc behind the dancer in order to obtain the dance information. The results suggest that bees are using the position of their own bodies to determine direction.     

East learns from West: Asiatic honeybees can understand dance language of European honeybees

The honeybee waggle dance, through which foragers advertise the existence and location of a food source to their hive mates, is acknowledged as the only known form of symbolic communication in an invertebrate. However, the suggestion, that different species of honeybee might possess distinct 'dialects' of the waggle dance, remains controversial. Furthermore, it remains unclear whether different species of honeybee can learn from and communicate with each other. This study reports experiments using a mixed-species colony that is composed of the Asiatic bee Apis cerana cerana (Acc), and the European bee Apis mellifera ligustica (Aml). Using video recordings made at an observation hive, we first confirm that Acc and Aml have significantly different dance dialects, even when made to forage in identical environments. When reared in the same colony, these two species are able to communicate with each other: Acc foragers could decode the dances of Aml to successfully locate an indicated food source. We believe that this is the first report of successful symbolic communication between two honeybee species; our study hints at the possibility of social learning between the two honeybee species, and at the existence of a learning component in the honeybee dance language.     

Dynamic range compression in the honey bee auditory system toward waggle dance sounds

Honey bee foragers use a "waggle dance" to inform nestmates about direction and distance to locations of attractive food. The sound and air flows generated by dancer's wing and abdominal vibrations have been implicated as important cues, but the decoding mechanisms for these dance messages are poorly understood. To understand the neural mechanisms of honey bee dance communication, we analyzed the anatomy of antenna and Johnston's organ (JO) in the pedicel of the antenna, as well as the mechanical and neural response characteristics of antenna and JO to acoustic stimuli, respectively. The honey bee JO consists of about 300-320 scolopidia connected with about 48 cuticular "knobs" around the circumference of the pedicel. Each scolopidium contains bipolar sensory neurons with both type I and II cilia. The mechanical sensitivities of the antennal flagellum are specifically high in response to low but not high intensity stimuli of 265-350 Hz frequencies. The structural characteristics of antenna but not JO neurons seem to be responsible for the non-linear responses of the flagellum in contrast to mosquito and fruit fly. The honey bee flagellum is a sensitive movement detector responding to 20 nm tip displacement, which is comparable to female mosquito. Furthermore, the JO neurons have the ability to preserve both frequency and temporal information of acoustic stimuli including the "waggle dance" sound. Intriguingly, the response of JO neurons was found to be age-dependent, demonstrating that the dance communication is only possible between aged foragers. These results suggest that the matured honey bee antennae and JO neurons are best tuned to detect 250-300 Hz sound generated during "waggle dance" from the distance in a dark hive, and that sufficient responses of the JO neurons are obtained by reducing the mechanical sensitivity of the flagellum in a near-field of dancer. This nonlinear effect brings about dynamic range compression in the honey bee auditory system.     

Does Imprecision in The Waggle Dance Fit Patterns Predicted by The Tuned-Error Hypothesis?

The waggle dance of the honey bee is used to recruit nest mates to a resource, though direction indicated for a resource may vary greatly within a single dance. Some authors suggest that this variation exits as an adaptation to distribute recruits across a patch of flowers, and that, due to the variation’s inverse relationship with distance, the shape of the recruit distribution will remain constant for resources at different distances. In this study, we test this hypothesis by examining how variation in the indication of direction and distance changes with respect to distance. We find that imprecision in the communication of direction does not diminish rapidly enough to accommodate an adaptive-error hypothesis, and we also find that variation in the indication of distance has a positive relationship with the distance of a resource from the hive.     

两种生境下野慈姑繁殖差异及其机制

植物的生长环境不但可直接作用于其可利用资源的多寡及其繁殖分配,而且可通过影响植物所在的群落以及传粉者的组成或行为而间接的导致繁殖差异。然而,直接与间接作用的方向或强度很少被同时关注,从而限制了从机制上理解环境对植物繁殖的影响。选取野慈姑(Sagittaria trifolia L.)为研究材料,将相同基因型组成的植株分别种植在光照差异明显的两个同质园(林荫区与日照区),于盛花期对野慈姑的开花数量、昆虫访花进行了观察,确定其主要访花昆虫、记录访花行为以及野慈姑的繁殖产出水平。研究表明,日照区野慈姑每天开放的花朵和植株数量显著高于林荫区。两个区域内野慈姑的主要访花者有四大类:蚜蝇类、蜂类、蝶类和蝇类;其中林荫区的主要传粉昆虫为蚜蝇类,而日照区为蜂类。日照区昆虫单位时间内的访花频率、昆虫每回合访问的雄花数、总花数和花序数均显著高于林荫区。日照区野慈姑的座果率显著高于林荫区,而单果种子数量与种子面积与林荫区相当。总体而言,日照区野慈姑的繁殖产出达到了林荫区的三倍以上,这是环境的直接作用与环境介导的传粉作用在同一方向上叠加的结果,且前者占主导地位。在关注植物、物理环境和生物因子三者之间互作的前提下,量化每个繁殖阶段的水平,特别是传粉昆虫在不同生境下的表现,阐述了环境异质性导致植物繁殖差异的根本原因。     

Increase in dance imprecision with decreasing foraging distance in the honey bee <Emphasis Type="Italic">Apis mellifera</Emphasis> L. is partly explained by physical constraints

Honey bee foragers communicate the direction and distance of both food sources and new nest sites to nest mates by means of a symbolic dance language. Interestingly, the precision by which dancers transfer directional information is negatively correlated with the distance to the advertised food source. The ‘tuned-error’ hypothesis suggests that colonies benefit from this imprecision as it spreads recruits out over a patch of constant size irrespective of the distance to the advertised site. An alternative to the tuned-error hypothesis is that dancers are physically incapable of dancing with great precision for nearby sources. Here we revisit the tuned-error hypothesis by studying the change in dance precision with increasing foraging distance over relatively short distances while controlling for environmental influences. We show that bees indeed increase their dance precision with the increase in foraging distance. However, we also show that dances performed by swarm-scouts for a nearby (30 m) nest site, where there could be no benefit to imprecision, are either without or with only limited directional information. This result suggests that imprecision in dance communication is caused primarily by physical constraints in the ability of dancers to turn around quickly enough when the advertised site is nearby.     

Responses to light under varying magnetic conditions in the honeybee,Apis mellifica

Summary In the dance of honeybees the indication of direction to a food source can be influenced by magnetic and photic stimuli. We have tested the behaviour of dancing honeybees illuminated with white light under varying magnetic conditions. The bees respond to the light stimulus with a maximum deviation from the correct dancing direction when they dance parallel to the inclination of the earth's magnetic field (EMF). The response to light drops to zero with increasing deviation from this zero-point direction (see also Martin and Lindauer 1977). The time of total indifference to light varies with the magnetic conditions. In the natural EMF the reaction to light becomes zero 20.3° (i.e. 1 h) after the bees have passed the zeropoint. In the compensated EMF this effect is delayed by 10°. The bees show nearly no reaction to light when the EMF is amplified to 2 Gauss.The relative spectral sensitivity of dancing honey-bees was tested in the compensated EMF. It is 1:1.63:2.64 for green-, blue-, and UV-light, respectively.Abbreviation EMF earth's magnetic field     

Colony migration in the tropical honey beeApis dorsata F. (Hymenoptera: Apidae)

Summary To obtain insights into the organization and adaptive significance of seasonal migration by colonies of the giant honey bee,Apis dorsata, we monitored the arrivals and departures of colonies in a rain forest habitat in northeastern Thailand, compared patterns of honey bee abundance with other measures of habitat variability, and observed the role of dance communication in organizing the migratory departure of a colony. Colonies arrived in the area during the end of the dry season, reproduced, and then departed early in the rainy season. During the immigration phase, early-arriving colonies stayed only temporarily, as if assessing habitat quality. Colonies departing after a long stay always left barren combs behind, suggesting that they had left in response to deteriorating resource quality. These observations support the idea that migration allows colonies to track seasonally varying resources in different regions. Our observations of a colony preparing for migration revealed that the dance language is involved in organizing the colony's departure, but that dancers signal only the direction to be taken, rather than, as in dances to feeding sites, both the direction and distance of a particular location.     

Mathematical analysis of the honeybee waggle dance

A honeybee informs her nestmates of the location of a flower by doing a waggle dance. The waggle dance encodes both the direction of and distance to the flower from the hive. To reveal how the waggle dance benefits the colony, we created a Markov model of bee foraging behavior and performed simulation experiments by incorporating the biological parameters that we obtained from our own observations of real bees as well as from the literature. When two feeders were each placed 400 m away from the hive in different directions, a virtual colony in which honeybees danced and correctly transferred information (a normal, real bee colony) made significantly greater numbers of successful visits to the feeders compared to a colony with inaccurate information transfer. Howerer, when five feeders were each located 400 m from the hive, the inaccurate information transfer colony performed better than the normal colony. These results suggest that dancing's ability to communicate accurate information depends on the number of feeders. Furthermore, because non-dancing colonies always made significantly fewer visits than those two colonies, we concluded that dancing behavior is beneficial for hives' ability to visit food sources.     

Interaction of gravi- and phototropic stimulation in the response of maize (Zea mays L.) coleoptiles

The influence of gravitropic stimulation upon blue-light-induced first positive phototropism for stimulations in the same (light source and center of gravity opposite to each other) and in opposing directions was investigated in maize cole-optiles by measuring fluence-response patterns. As a result of gravitropic counterstimulation, phototropic bending was transient with maximum curvature occurring 100 min after stimulation. On a horizontal clinostat, however, the seedlings curved for 20 h. Gravistimulation in the opposite direction acted additively upon blue-light curvature. Gravistimulation in the same direction as phototropic stimulation produced a complex behaviour deviating from simple additivity. This pattern can be explained by a gravitropically mediated sensitization of the phototropic reaction, an optimal dependence of differential growth on the sum of photo-and gravistimulation, and blue-light-induced inhibition of gravitropic curvature at high fluences. These findings indicate that several steps of photo-and gravitransduction are separate. Preirradiation with red light desensitized the system independently of applied gravity-treatment, indicating that the site of red-light interaction is common to both transduction chains.Abbreviations BL blue light - G⁺ stimulation by light and gravity in the same direction (i.e. light source and center of gravity opposite to each other) - G^- stimulation by light and gravity in opposing directions     

Informational conflicts created by the waggle dance

The honeybee (Apis mellifera) waggle dance is one of the most intriguing animal communication signals. A dancing bee communicates the location of a profitable food source and its odour. Followers may often experience situations in which dancers indicate an unfamiliar location but carry the scent of a flower species the followers experienced previously at different locations. Food scents often reactivate bees to resume food collection at previously visited food patches. This double function of the dance creates a conflict between the social vector information and the private navigational information. We investigated which kind of information followers with field experience use in this situation and found that followers usually ignored the spatial information encoded by the waggle dance even if they followed a dance thoroughly (five waggle runs or more). They relied on private information about food source locations instead (in 93% of all cases). Furthermore, foragers preferred to follow dancers carrying food odours they knew from previous field trips, independently of the spatial information encoded in the dance. Surprisingly, neither odour identity nor the location indicated by the dancer was an important factor for the reactivation success of a dance. Our results contrast with the assumption that (i) followers usually try to decode the vector information and (ii) dances indicating an unfamiliar location are of little interest to experienced foragers.     

Not looking where you are leaping: a novel method of oriented travel in the caterpillar Calindoea trifascialis (Moore) (Lepidoptera: Thyrididae)

The prepupation caterpillar of the Southeast Asian moth Calindoea trifascialis constructs a leaf shelter that jumps across the ground using a jumping method novel among the insects. We found that movement path direction was correlated to the direction opposite to the most intense light. Correlated random walk (CRW) analyses found net squared displacements higher than predicted by a CRW, and fractal dimension analysis indicated straighter paths at large spatial scales. Rearing experiments showed high mortality from predation on the ground, but higher mortality resulted from sun exposure. We interpret jumping path orientation as an efficient search strategy to find shade in a variable landscape, given limited perception, in the presence of overheating and desiccation risks.     

Honeybee navigation: critically examining the role of the polarization compass

Although it is widely accepted that honeybees use the polarized-light pattern of the sky as a compass for navigation, there is little direct evidence that this information is actually sensed during flight. Here, we ask whether flying bees can obtain compass cues derived purely from polarized light, and communicate this information to their nest-mates through the ‘waggle dance’. Bees, from an observation hive with vertically oriented honeycombs, were trained to fly to a food source at the end of a tunnel, which provided overhead illumination that was polarized either parallel to the axis of the tunnel, or perpendicular to it. When the illumination was transversely polarized, bees danced in a predominantly vertical direction with waggles occurring equally frequently in the upward or the downward direction. They were thus using the polarized-light information to signal the two possible directions in which they could have flown in natural outdoor flight: either directly towards the sun, or directly away from it. When the illumination was axially polarized, the bees danced in a predominantly horizontal direction with waggles directed either to the left or the right, indicating that they could have flown in an azimuthal direction that was 90° to the right or to the left of the sun, respectively. When the first half of the tunnel provided axial illumination and the second half transverse illumination, bees danced along all of the four principal diagonal directions, which represent four equally likely locations of the food source based on the polarized-light information that they had acquired during their journey. We conclude that flying bees are capable of obtaining and signalling compass information that is derived purely from polarized light. Furthermore, they deal with the directional ambiguity that is inherent in polarized light by signalling all of the possible locations of the food source in their dances, thus maximizing the chances of recruitment to it.     

Honey bee waggle dance communication increases diversity of pollen diets in intensively managed agricultural landscapes

The benefits of honey bee dance communication for colony performance in different resource environments are still not well understood. Here, we test the hypothesis that directional dance communication enables honey bee colonies to maintain a diverse pollen diet, especially in landscapes with low resource diversity. To test this hypothesis, we placed 24 Apis mellifera L. colonies with either intact or experimentally disrupted dance communication in eight agricultural landscapes that differed in the diversity of flowering plants and in the dominance of mass‐flowering crops. Pollen from incoming foragers was collected and identified via DNA metabarcoding. Disrupting dance communication affected the way the diversity of honey bee pollen diets was impacted by the dominance of mass‐flowering crops in available flower resources (p = .04). With increasing dominance of mass‐flowering crops in resource environments, foragers of colonies with intact communication foraged on an increasing proportion of available plant genera (p = .01). This was not the case for colonies with disrupted dance communication (p = .5). We conclude that the honey bee dance communication benefits pollen foraging on diverse plant resources and thereby contributes to high quality nutrition in environments with low‐resource diversity.