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In this work, we identified the trail pheromone of the ant Crematogaster scutellaris. We combined gas chromatography-mass spectrometry analysis of extracts from the hind tibia, the location of the respective glands, with automated trail following assays. The study found tridecan-2-ol to be the strongest discriminator between hind tibia and other body part extracts. Tridecan-2-ol elicited trail-following behaviour at concentrations of 1 ng/µL. A separation of the enantiomers showed responses to (R)-tridecan-2-ol already at 0.001 ng/µL and only at a 1000-fold higher concentration for (S)-tridecan-2-ol, suggesting that only the R enantiomer is used by C. scutellaris in its natural environment. We also found strong behavioural responses to 2-dodecanol, a substance that was not detectable in the hind tibia extract of C. scutellaris, but which has been reported to be the trail pheromone of the related species C. castanea. We discuss the contribution of these results to the 'dissection and reconstruction' of strategies and mechanisms underlying the social organization of ants.
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Hormigas , Feromonas , Animales , Feromonas/análisis , Hormigas/fisiología , Conducta Animal , Conducta AlimentariaRESUMEN
Geosmin is an odorant produced by bacteria in moist soil. It has been found to be extraordinarily relevant to some insects, but the reasons for this are not yet fully understood. Here we report the first tests of the effect of geosmin on honey bees. A stinging assay showed that the defensive behaviour elicited by the bee's alarm pheromone component isoamyl acetate (IAA) is strongly suppressed by geosmin. Surprisingly, the suppression is, however, only present at very low geosmin concentrations, and disappears at higher concentrations. We investigated the underlying mechanisms at the level of the olfactory receptor neurons by means of electroantennography, finding the responses to mixtures of geosmin and IAA to be lower than to pure IAA, suggesting an interaction of both compounds at the olfactory receptor level. Calcium imaging of the antennal lobe (AL) revealed that neuronal responses to geosmin decreased with increasing concentration, correlating well with the observed behaviour. Computational modelling of odour transduction and coding in the AL suggests that a broader activation of olfactory receptor types by geosmin in combination with lateral inhibition could lead to the observed non-monotonic increasing-decreasing responses to geosmin and thus underlie the specificity of the behavioural response to low geosmin concentrations.
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Receptores Odorantes , Abejas , Animales , Odorantes , Feromonas/farmacología , NaftolesRESUMEN
All animals need information about the direction of motion to be able to track the trajectory of a target (prey, predator, cospecific) or to control the course of navigation. This information is provided by direction selective (DS) neurons, which respond to images moving in a unique direction. DS neurons have been described in numerous species including many arthropods. In these animals, the majority of the studies have focused on DS neurons dedicated to processing the optic flow generated during navigation. In contrast, only a few studies were performed on DS neurons related to object motion processing. The crab Neohelice is an established experimental model for the study of neurons involved in visually-guided behaviors. Here, we describe in male crabs of this species a new group of DS neurons that are highly directionally selective to moving objects. The neurons were physiologically and morphologically characterized by intracellular recording and staining in the optic lobe of intact animals. Because of their arborization in the lobula complex, we called these cells lobula complex directional cells (LCDCs). LCDCs also arborize in a previously undescribed small neuropil of the lateral protocerebrum. LCDCs are responsive only to horizontal motion. This nicely fits in the behavioral adaptations of a crab inhabiting a flat, densely crowded environment, where most object motions are generated by neighboring crabs moving along the horizontal plane.SIGNIFICANCE STATEMENT Direction selective (DS) neurons are key to a variety of visual behaviors including, target tracking (preys, predators, cospecifics) and course control. Here, we describe the physiology and morphology of a new group of remarkably directional neurons exclusively responsive to horizontal motion in crabs. These neurons arborize in the lobula complex and in a previously undescribed small neuropil of the lateral protocerebrum. The strong sensitivity of these cells for horizontal motion represents a clear example of functional neuronal adaptation to the lifestyle of an animal inhabiting a flat environment.
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Adaptación Fisiológica , Braquiuros/fisiología , Percepción de Movimiento/fisiología , Movimiento , Neuronas/fisiología , Potenciales de Acción , Animales , Braquiuros/citología , Masculino , Neuronas/citología , Lóbulo Óptico de Animales no Mamíferos/citología , Lóbulo Óptico de Animales no Mamíferos/fisiologíaRESUMEN
Animals use binocular information to guide many behaviors. In highly visual arthropods, complex binocular computations involved in processing panoramic optic flow generated during self-motion occur in the optic neuropils. However, the extent to which binocular processing of object motion occurs in these neuropils remains unknown. We investigated this in a crab, where the distance between the eyes and the extensive overlapping of their visual fields advocate for the use of binocular processing. By performing in vivo intracellular recordings from the lobula (third optic neuropil) of male crabs, we assessed responses of object-motion-sensitive neurons to ipsilateral or contralateral moving objects under binocular and monocular conditions. Most recorded neurons responded to stimuli seen independently with either eye, proving that each lobula receives profuse visual information from both eyes. The contribution of each eye to the binocular response varies among neurons, from those receiving comparable inputs from both eyes to those with mainly ipsilateral or contralateral components, some including contralateral inhibition. Electrophysiological profiles indicated that a similar number of neurons were recorded from their input or their output side. In monocular conditions, the first group showed shorter response delays to ipsilateral than to contralateral stimulation, whereas the second group showed the opposite. These results fit well with neurons conveying centripetal and centrifugal information from and toward the lobula, respectively. Intracellular and massive stainings provided anatomical support for this and for direct connections between the two lobulae, but simultaneous recordings failed to reveal such connections. Simplified model circuits of interocular connections are discussed.SIGNIFICANCE STATEMENT Most active animals became equipped with two eyes, which contributes to functions like depth perception, objects spatial location, and motion processing, all used for guiding behaviors. In visually active arthropods, binocular neural processing of the panoramic optic flow generated during self-motion happens already in the optic neuropils. However, whether binocular processing of single-object motion occurs in these neuropils remained unknown. We investigated this in a crab, where motion-sensitive neurons from the lobula can be recorded in the intact animal. Here we demonstrate that different classes of neurons from the lobula compute binocular information. Our results provide new insight into where and how the visual information acquired by the two eyes is first combined in the brain of an arthropod.
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Braquiuros/fisiología , Percepción de Movimiento/fisiología , Neuronas/fisiología , Neurópilo/fisiología , Visión Binocular/fisiología , Vías Visuales/fisiología , Animales , Dominancia Cerebral , Masculino , Neurópilo/ultraestructura , Técnicas de Placa-Clamp , Estimulación Luminosa , Tiempo de Reacción , Visión Monocular/fisiologíaRESUMEN
Highly active insects and crabs depend on visual motion information for detecting and tracking mates, prey, or predators, for which they require directional control systems containing internal maps of visual space. A neural map formed by large, motion-sensitive neurons implicated in processing panoramic flow is known to exist in an optic ganglion of the fly. However, an equivalent map for processing spatial positions of single objects has not been hitherto identified in any arthropod. Crabs can escape directly away from a visual threat wherever the stimulus is located in the 360° field of view. When tested in a walking simulator, the crab Neohelice granulata immediately adjusts its running direction after changes in the position of the visual danger stimulus smaller than 1°. Combining mass and single-cell staining with in vivo intracellular recording, we show that a particular class of motion-sensitive neurons of the crab's lobula that project to the midbrain, the monostratified lobula giants type 1 (MLG1), form a system of 16 retinotopically organized elements that map the 360° azimuthal space. The preference of these neurons for horizontally moving objects conforms the visual ecology of the crab's mudflat world. With a mean receptive field of 118°, MLG1s have a large superposition among neighboring elements. Our results suggest that the MLG1 system conveys information on object position as a population vector. Such computational code can enable the accurate directional control observed in the visually guided behaviors of crabs.
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Braquiuros/fisiología , Percepción de Movimiento/fisiología , Red Nerviosa/fisiología , Lóbulo Óptico de Animales no Mamíferos/citología , Orientación/fisiología , Células Receptoras Sensoriales/fisiología , Vías Visuales/fisiología , Potenciales de Acción/fisiología , Animales , Braquiuros/citología , Reacción de Fuga/fisiología , Técnicas In Vitro , Masculino , Movimiento/fisiología , Distribución Normal , Campos Visuales/fisiologíaRESUMEN
Historically, arthropod behavior has been considered to be a collection of simple, automaton-like routines commanded by domain-specific brain modules working independently. Nowadays, it is evident that the extensive behavioral repertoire of these animals and its flexibility necessarily imply far more complex abilities than originally assumed. For example, even what was thought to be a straightforward behavior of crabs, the escape response to visual danger stimuli, proved to involve a number of sequential stages, each of which implying decisions made on the bases of stimulus and contextual information. Inspired in previous observations on how the stimulus trajectory can affect the escape response of crabs in the field, we investigated the escape response to images of objects approaching directly toward the crab (looming stimuli: LS) or moving parallel to it (translational stimuli: TS) in the laboratory. Computer simulations of moving objects were effective to elicit escapes. LS evoked escapes with higher probability and intensity (speed and distance of escape) than TS, but responses started later. In addition to the escape run, TS also evoked a defensive response of the animal with its claws. Repeated presentations of TS or LS were both capable of inducing habituation. Results are discussed in connection with the possibilities offered by crabs to investigate the neural bases of behaviors occurring in the natural environment.