Effect of juvenile hormone on short-term olfactory memory in young honeybees (Apis mellifera).

Reliable retention of olfactory learning following a 1-trial classical conditioning of the proboscis extension reflex (PER) is not achieved in honeybees until they are 6-7 days old. Here we show that treatment of newly emerged honeybees with juvenile hormone (JH) has a profound effect on the maturation of short-term olfactory memory. JH-treated individuals display excellent short-term (1 h) memory of associative learning at times as early as 3 days of age and perform consistently better than untreated bees for at least the first week of their lives. By contrast, the retention of long-term (24 h) memory following a 3-trial conditioning of the PER is not significantly improved in JH-treated bees. Our study also shows that experience and (or) chemosensory activation are not essential to improve learning performance in olfactory tasks. The lack of accelerated development of long-term retention of olfactory memories in JH-treated honeybees is discussed in the context of neural circuits suspected to mediate memory formation and retrieval in the honeybee brain.     

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.     

蜜蜂对迷宫的学习和记忆

著名的生物学家 Frisch[1] 研究蜜蜂的行为时发现 ,蜜蜂可以从所有其它颜色中区别蓝色 ,说明蜜蜂能够看到作为一种颜色的蓝色 ,并能学会以颜色作为食物的信号。从此 ,揭开了研究蜜蜂学习和记忆的序幕 ,Menzel[2 ] 把蜜蜂作为一个模型系统来研究学习与记忆。长期以来 ,许多生物学家的大量研究发现 :蜜蜂不仅对目标的颜色、模式、气味等有较好的学习和记忆能力 ,而且对陆标以及时间模式等都有较好的学习和记忆能力。然而 ,研究蜜蜂学习迷宫的工作相对比较少 ,Weiss[3] 训练蜜蜂步行通过了相对简单的迷宫 ,Menze[4 ] 在一个 T型迷宫中训练步…     

Response of olfactory receptor neurons in honeybees to odorants and their binary mixtures

Recordings were made from single sensilla placodea of the worker honeybee (Apis mellifera). The sensilla were stimulated with one of two sets of four compounds and their binary mixtures, at two dosage levels. Aromatic compounds comprised one set, and saturated n-octane derivatives comprised the other set. Correlation, principal component, and cluster analyses indicate that responses to binary mixtures are not linear combinations of responses to the component compounds. The first principal component indicated that neuronal units had either more excitatory or more inhibitory responses to all odorants than would be expected from a model where inhibitory and excitatory responses are randomly distributed among the neuronal units. When compared to the responses to the component odorants, synergistic responses to binary odors occurred more often than would be expected by chance. Clear inhibitory responses to binary odors were less prevalent. This study agrees with an earlier study employing aromatic odorants in that most of the aromatic odorants each had groups of receptor neurons that were relatively selective for it, and each odorant had a distinctly different number of receptor neurons selective for it. Among the octane derivatives, receptor neurons were selective for the level of oxidation of the functional group or its site of attachment, rather than specific compounds.     

Visual generalization in honeybees: evidence of peak shift in color discrimination

In the present study, we investigated color generalization in the honeybee Apis mellifera after differential conditioning. In particular, we evaluated the effect of varying the position of a novel color along a perceptual continuum relative to familiar colors on response biases. Honeybee foragers were differentially trained to discriminate between rewarded (S+) and unrewarded (S?) colors and tested on responses toward the former S+ when presented against a novel color. A color space based on the receptor noise-limited model was used to evaluate the relationship between colors and to characterize a perceptual continuum. When S+ was tested against a novel color occupying a locus in the color space located in the same direction from S? as S+, but further away, the bees shifted their stronger response away from S? toward the novel color. These results reveal the occurrence of peak shift in the color vision of honeybees and indicate that honeybees can learn color stimuli in relational terms based on chromatic perceptual differences.     

Pollinator shift to managed honeybees enhances reproductive output in a bumblebee-pollinated plant

Introduced honeybees have had a large impact on native ecosystems by disrupting native plant–pollinator interactions. However, little is known of the effect of honeybees on reproduction of bumblebee-pollinated plants. Seasonal displacement of native bumblebees by introduced honeybees (Apis mellifera and A. cerana) was observed in Pedicularis densispica, endemic to Hengduan Mountains, China, providing an opportunity for honeybee presence/absence comparisons. Five-year field surveys were conducted in one frequently disturbed population at Yila Pasture (YP). We compared pollination effectiveness (combinations of visitation rate, efficiency in pollen transfer, and potential geitonogamy) between native and introduced managed bees. The total visitation rate of native bees and subsequent reproductive output decreased progressively, but honeybee introduction resulted in at least twofold increase in visitation and 70 % increase in seed set. In general, native bumblebees, which have larger bodies and longer proboscises and spent more time probing single flowers, were more efficient than honeybees in terms of pollen removal and pollen deposition during first visits to virgin flowers. Compared with bumblebees, honeybees visited markedly fewer flowers in sequence within individual plants, potentially reducing geitonogamous pollination. Our data highlight that introduced honeybees can provide pollination service in terms of both quantity and quality for P. densispica. We suggest honeybee introduction as an effective way to augment pollination of P. densispica at disturbed and isolated sites.     

Drosophila olfactory memory: single genes to complex neural circuits

A central goal of neuroscience is to understand how neural circuits encode memory and guide behaviour. Studying simple, genetically tractable organisms, such as Drosophila melanogaster, can illuminate principles of neural circuit organization and function. Early genetic dissection of D. melanogaster olfactory memory focused on individual genes and molecules. These molecular tags subsequently revealed key neural circuits for memory. Recent advances in genetic technology have allowed us to manipulate and observe activity in these circuits, and even individual neurons, in live animals. The studies have transformed D. melanogaster from a useful organism for gene discovery to an ideal model to understand neural circuit function in memory.     

Aversive learning in honeybees revealed by the olfactory conditioning of the sting extension reflex

Invertebrates have contributed greatly to our understanding of associative learning because they allow learning protocols to be combined with experimental access to the nervous system. The honeybee Apis mellifera constitutes a standard model for the study of appetitive learning and memory since it was shown, almost a century ago, that bees learn to associate different sensory cues with a reward of sugar solution. However, up to now, no study has explored aversive learning in bees in such a way that simultaneous access to its neural bases is granted. Using odorants paired with electric shocks, we conditioned the sting extension reflex, which is exhibited by harnessed bees when subjected to a noxious stimulation. We show that this response can be conditioned so that bees learn to extend their sting in response to the odorant previously punished. Bees also learn to extend the proboscis to one odorant paired with sugar solution and the sting to a different odorant paired with electric shock, thus showing that they can master both appetitive and aversive associations simultaneously. Responding to the appropriate odorant with the appropriate response is possible because two different biogenic amines, octopamine and dopamine subserve appetitive and aversive reinforcement, respectively. While octopamine has been previously shown to substitute for appetitive reinforcement, we demonstrate that blocking of dopaminergic, but not octopaminergic, receptors suppresses aversive learning. Therefore, aversive learning in honeybees can now be accessed both at the behavioral and neural levels, thus opening new research avenues for understanding basic mechanisms of learning and memory.     

From honeybees to Internet servers: biomimicry for distributed management of Internet hosting centers

An Internet hosting center hosts services on its server ensemble. The center must allocate servers dynamically amongst services to maximize revenue earned from hosting fees. The finite server ensemble, unpredictable request arrival behavior and server reallocation cost make server allocation optimization difficult. Server allocation closely resembles honeybee forager allocation amongst flower patches to optimize nectar influx. The resemblance inspires a honeybee biomimetic algorithm. This paper describes details of the honeybee self-organizing model in terms of information flow and feedback, analyzes the homology between the two problems and derives the resulting biomimetic algorithm for hosting centers. The algorithm is assessed for effectiveness and adaptiveness by comparative testing against benchmark and conventional algorithms. Computational results indicate that the new algorithm is highly adaptive to widely varying external environments and quite competitive against benchmark assessment algorithms. Other swarm intelligence applications are briefly surveyed, and some general speculations are offered regarding their various degrees of success.     

Ceramide: From lateral segregation to mechanical stress

Ceramide is a sphingolipid present in eukaryotic cells that laterally segregates into solid domains in model lipid membranes. Imaging has provided a wealth of structural information useful to understand some of the physical properties of these domains. In biological membranes, ceramide is formed on one of the membrane leaflets by enzymatic cleavage of sphyngomyelin. Ceramide, with a smaller head size than its parent compound sphyngomyelin, induces an asymmetric membrane tension and segregates into highly ordered domains that have a much high shear viscosity than that of the surrounding lipids. These physical properties, together with the rapid transmembrane flip-flop of the locally produced ceramide, trigger a sequence of membrane perturbations that could explain the molecular mechanism by which ceramide mediates different cell responses. In this review we will try to establish a connection between the physical membrane transformations in model systems known to occur upon ceramide formation and some physiologically relevant process in which ceramide is known to participate.     

Cutting edge: local recall responses by memory T cells newly recruited to peripheral nonlymphoid tissues

Infection results in the formation of a circulating effector memory T cell population able to enter peripheral tissues either in the steady state or in response to localized infection. As a consequence, recall is thought to result from a phased response first involving those T cells already at the site of infection followed by the infiltration of memory cells from the wider circulation. We have recently reported that tissue-resident T cells can undergo stimulation and proliferation in response to local infection. In this study, we examine the proliferation of memory T cells newly recruited from the circulation. Our results show that although recruitment of circulating memory cells is nonspecific in nature, there is preferential proliferation of specific T cells within infected tissues. Thus, expansion represents a means of local Ag-specific enrichment of T cells recruited from a circulating memory pool of mixed specificities.     

Reward quality influences the development of learned olfactory biases in honeybees

Plants produce flowers with complex visual and olfactory signals, but we know relatively little about the way that signals such as floral scents have evolved. One important factor that may direct the evolution of floral signals is a pollinator''s ability to learn. When animals learn to associate two similar signals with different outcomes, biases in their responses to new signals can be formed. Here, we investigated whether or not pollinators develop learned biases towards floral scents that depend on nectar reward quality by training restrained honeybees to learn to associate two similar odour signals with different outcomes using a classical conditioning assay. Honeybees developed learned biases towards odours as a result of differential conditioning, and the extent to which an olfactory bias could be produced depended upon the difference in the quality of the nectar rewards experienced during conditioning. Our results suggest that differences in reward quality offered by flowers influence odour recognition by pollinators, which in turn could influence the evolution of floral scents in natural populations of co-flowering plants.     

Behavioral studies on tarsal gustation in honeybees: sucrose responsiveness and sucrose-mediated olfactory conditioning

Although the forelegs of honeybees are one of their main gustatory appendages, tarsal gustation in bees has never been systematically studied. To provide a more extensive account on honeybee tarsal gustation, we performed a series of behavioral experiments aimed at characterizing (1) tarsal sucrose sensitivity under different experimental conditions and (2) the capacity of tarsal sucrose stimulation to support olfactory conditioning. We quantified the proboscis extension reflex to tarsal sucrose stimulation and to odors paired with tarsal sucrose stimulation, respectively. Our experiments show that tarsal sucrose sensitivity is lower than antennal sucrose sensitivity and can be increased by starvation time. In contrast, antennae amputation decreases tarsal sucrose sensitivity. Furthermore, we show that tarsal sucrose stimulation can support olfactory learning and memory even if the acquisition level reached is relatively low (40%).     

Secret life of plants: From memory to intelligence

Plants are able to perform photosynthesis and cannot escape from environmental stresses, so they therefore developed sophisticated, highly responsive and dynamic physiology. Others'' and our results indicate that plants solve their optimal light acclimation and immune defenses, photosynthesis and transpiration by a computational algorithm of the cellular automation. Our recent results however suggest that plants are capable of processing information encrypted in light intensity and in its energy. With the help of nonphotochemical quenching and photoelectrophysiological signaling (PEPS) plants are able to perform biological quantum computation and memorize light training in order to optimize their Darwinian fitness. Animals have their network of neuron synapses, electrophysiological circuits and memory, but plants have their network of chloroplasts connected by stromules, PEPS circuits transduced by bundle sheath cells and cellular light memory. It is suggested that plants could be intelligent organisms with much higher organism organization levels than it was thought before.Key words: excess excitation energy, cellular automation, cellular light memory, Darwinian fitness, nonphotochemical quenching, photoelectrophysiological signaling, SAA, SAR