RESUMEN
Spiders show a broad range of motions in addition to walking and running with their eight coordinated legs taking them towards their resources and away from danger. The usefulness of all these motions depends on the ability to control and adjust them to changing environmental conditions. A remarkable wealth of sensory receptors guarantees the necessary guidance. Many facets of such guidance have emerged from neuroethological research on the wandering spider Cupiennius salei and its allies, although sensori-motor control was not the main focus of this work. The present review may serve as a springboard for future studies aiming towards a more complete understanding of the spider's control of its different types of motion. Among the topics shortly addressed are the involvement of lyriform slit sensilla in path integration, muscle reflexes in the walking legs, the monitoring of joint movement, the neuromuscular control of body raising, the generation of vibratory courtship signals, the sensory guidance of the jump to flying prey and the triggering of spiderling dispersal behavior. Finally, the interaction of sensors on different legs in oriented turning behavior and that of the sensory systems for substrate vibration and medium flow are addressed.
Asunto(s)
Locomoción/fisiología , Desempeño Psicomotor/fisiología , Sensilos/fisiología , Arañas/fisiología , Animales , Mecanorreceptores/fisiología , Movimiento/fisiología , Órganos de los Sentidos/fisiología , Células Receptoras Sensoriales/fisiologíaRESUMEN
The measurement of cuticular strain during locomotion using foil strain gauges provides information both on the loads of the exoskeleton bears and the adaptive value of the specific location of natural strain detectors (slit sense organs). Here, we critically review available literature. In tethered animals, by applying loads to the metatarsus tip, strain and mechanical sensitivity (S = strain/load) induced at various sites in the tibia were determined. The loci of the lyriform organs close to the tibia-metatarsus joint did not stand out by high strain. The strains induced at various sites during free locomotion can be interpreted based on S and, beyond the joint region, on beam theory. Spiders avoided laterad loading of the tibia-metatarsus joint during slow locomotion. Balancing body weight, joint flexors caused compressive strain at the posterior and dorsal tibia. While climbing upside down strain measurements indicate strong flexor activity. In future studies, a precise calculation and quantitative determination of strain at the sites of the lyriform organs will profit from more detailed data on the overall strain distribution, morphology, and material properties. The values and caveats of the strain gauge technology, the only one applicable to freely moving spiders, are discussed.
Asunto(s)
Exoesqueleto/fisiología , Fenómenos Biomecánicos/fisiología , Extremidades/fisiología , Locomoción/fisiología , Arañas/fisiología , Estrés Mecánico , Animales , VirtudesRESUMEN
The work of the Nobel Laureate Karl von Frisch, the founder of this journal, was seminal in many ways. He established the honeybee as a key animal model for experimental behavioural studies on sensory perception, learning and memory, and first correctly interpreted its famous dance communication. Here, we report on a previously unknown letter by the Physicist and Nobel Laureate Albert Einstein that was written in October 1949. It briefly addresses the work of von Frisch and also queries how understanding animal perception and navigation may lead to innovations in physics. We discuss records proving that Einstein and von Frisch met in April 1949 when von Frisch visited the USA to present a lecture on bees at Princeton University. In the historical context of Einstein's theories and thought experiments, we discuss some more recent discoveries of animal sensory capabilities alien to us humans and potentially valuable for bio-inspired design improvements. We also address the orientation of animals like migratory birds mentioned by Einstein 70 years ago, which pushes the boundaries of our understanding nature, both its biology and physics.
Asunto(s)
Abejas , Conducta Animal , Correspondencia como Asunto/historia , Animales , Historia del Siglo XX , HumanosRESUMEN
Engineered systems are typically based on a large variety of materials differing in composition and processing to provide the desired functionality. Nature, however, has evolved materials that are used for a wide range of functional challenges with minimal compositional changes. The exoskeletal cuticle of spiders, as well as of other arthropods such as insects and crustaceans, is based on a combination of chitin, protein, water and small amounts of organic cross-linkers or minerals. Spiders use it to obtain mechanical support structures and lever systems for locomotion, protection from adverse environmental influences, tools for piercing, cutting and interlocking, auxiliary structures for the transmission and filtering of sensory information, structural colours, transparent lenses for light manipulation and more. This paper illustrates the 'design space' of a single type of composite with varying internal architecture and its remarkable capability to serve a diversity of functions. This article is part of the theme issue 'Bio-derived and bioinspired sustainable advanced materials for emerging technologies (part 1)'.
Asunto(s)
Arañas , Animales , Quitina , Crustáceos , Minerales , ProteínasRESUMEN
Non-nervous auxiliary structures play a significant role in sensory biology. They filter the stimulus and transform it in a way that fits the animal's needs, thereby contributing to the avoidance of the central nervous system's overload with meaningless stimuli and a corresponding processing task. The present review deals with mechanoreceptors mainly of invertebrates and some remarkable recent findings stressing the role of mechanics as an important source of sensor adaptedness, outstanding performance, and diversity. Instead of organizing the review along the types of stimulus energy (force) taken up by the sensors, processes associated with a few basic and seemingly simple mechanical principles like lever systems, viscoelasticity, resonance, traveling waves, and impedance matching are taken as the guideline. As will be seen, nature makes surprisingly competent use of such "simple mechanics".
Asunto(s)
Fenómenos Biomecánicos/fisiología , Mecanorreceptores/fisiología , Mecanotransducción Celular/fisiología , AnimalesRESUMEN
Synchrotron X-ray diffraction was applied to study the structure of biogenic α-chitin crystals composing the tendon of the spider Cupiennius salei. Measurements were carried out on pristine chitin crystals stabilized by proteins and water, as well as after their deproteinization and dehydration. We found substantial shifts (up to Δq/q=9% in the wave vector in q-space) in the (020) diffraction peak position between intact and purified chitin samples. However, chitin lattice parameters extracted from the set of reflections (hkl), which did not contain the (020)-reflection, showed no systematic variation between the pristine and the processed samples. The observed shifts in the (020) peak position are discussed in terms of the ordering-induced modulation of the protein and water electron density near the surface of the ultra-thin chitin fibrils due to strong protein/chitin and water/chitin interactions. The extracted modulation periods can be used as a quantitative parameter characterizing the interaction length.
Asunto(s)
Quitina/química , Proteínas de Insectos/química , Nanopartículas/química , Animales , Modelos Químicos , Dispersión de Radiación , Espectrometría Raman , Arañas , Tendones/química , Agua/química , Difracción de Rayos XRESUMEN
Stingless bees (Hymenoptera, Apidae, Meliponini) are by far the largest group of eusocial bees on Earth. Due to the diversity of evolutionary responses to specific ecological challenges, the Meliponini are well suited for comparative studies of the various adaptations to the environment found in highly eusocial bees. Of particular interest are the physiological mechanisms underlying the sophisticated cooperative and collective actions of entire colonies, which form the basis of the ecological success of the different bee species under the particular conditions prevailing in their respective environment. The present Special Issue of the Journal of Comparative Physiology A provides a sample of the exciting diversity of sensorial and behavioral adaptations in stingless bees, particularly concerning (1) the sensory bases for foraging, (2) chemical communication, and (3) the behavioral ecology of foraging.
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Abejas/fisiología , Conducta Animal/fisiología , Sensación/fisiología , Animales , AmbienteRESUMEN
In stingless bees (Meliponini) like in many other eusocial insect colonies food hoarding plays an important role in colony survival. However, very little is known on how Meliponini, a taxon restricted to tropical and subtropical regions, respond to different store conditions. We studied the impact of honey removal on nectar foraging activity and recruitment behaviour in Melipona scutellaris and compared our results with studies of the honey bee Apis mellifera. As expected, foraging activity increased significantly during abundance of artificial nectar and when increasing its profitability. Foraging activity on colony level could thereby frequently increase by an order of magnitude. Intriguingly, however, poor honey store conditions did not induce increased nectar foraging or recruitment activity. We discuss possible reasons explaining why increasing recruitment and foraging activity are not used by meliponines to compensate for poor food conditions in the nest. Among these are meliponine specific adaptations to climatic and environmental conditions, as well as physiology and brood rearing, such as mass provisioning of the brood.
Asunto(s)
Abejas/fisiología , Conducta Alimentaria/fisiología , Privación de Alimentos/fisiología , Miel , Néctar de las Plantas/administración & dosificación , AnimalesRESUMEN
Adding to previous efforts towards a better understanding of the remarkable diversity of spider mechanosensitive hair sensilla, this study examines hairs of Cupiennius salei most likely serving a proprioreceptive function. At the tibia-metatarsus joint of all walking legs, there are two opposing groups of hairs ventrally on the tibia (20 hairs) and metatarsus (75 hairs), respectively. These hairs deflect each other when the joint flexes during locomotion, reversibly interlocking by microtrichs on their hair shafts. The torque resisting the hair deflection into the direction of natural stimulation is smaller by up to two powers of ten than that for the other directions. The torsional restoring constant S of the hair suspension is about 10(-10) Nm rad(-1) in the preferred direction, up to a hair deflection angle of 30° (mean of natural deflection angles). Joint movements were imposed in ranges and at rates measured in walking spiders and sensory action potentials recorded. Within the natural step frequencies (0.3-3 Hz) the rate of action potentials follows the velocity of hair deflection. All findings point to the morphological, mechanical, and physiological adaptedness of the joint hair sensilla to their proprioreceptive stimulation during locomotion.
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Adaptación Fisiológica , Locomoción/fisiología , Propiocepción/fisiología , Sensilos/fisiología , Arañas/fisiología , Potenciales de Acción , Animales , Fenómenos Biomecánicos , Articulaciones/fisiología , Microelectrodos , Microscopía Electrónica de Rastreo , Estimulación Física , Sensilos/ultraestructura , Umbral Sensorial , Arañas/ultraestructura , TorqueRESUMEN
Living organisms use composite materials for various functions, such as mechanical support, protection, motility and the sensing of signals. Although the individual components of these materials may have poor mechanical qualities, they form composites of polymers and minerals with a remarkable variety of functional properties. Researchers are now using these natural systems as models for artificial mechanosensors and actuators, through studying both natural structures and their interactions with the environment. In addition to inspiring the design of new materials, analysis of natural structures on this basis can provide insight into evolutionary constraints on structure-function relationships in living organisms and the variety of structural solutions that emerged from these constraints.
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Materiales Biocompatibles/metabolismo , Modelos Biológicos , Fenómenos Fisiológicos de las Plantas , Arañas/fisiología , Animales , Fenómenos Biomecánicos , Humanos , Mecanorreceptores/fisiología , Arañas/anatomía & histologíaRESUMEN
In contrast to marking of the location of resources or sexual partners using single-spot pheromone sources, pheromone paths attached to the substrate and assisting orientation are rarely found among flying organisms. However, they do exist in meliponine bees (Apidae, Apinae, Meliponini), commonly known as stingless bees, which represent a group of important pollinators in tropical forests. Worker bees of several Neotropical meliponine species, especially in the genus Scaptotrigona Moure 1942, deposit pheromone paths on substrates between highly profitable resources and their nest. In contrast to past results and claims, we find that these pheromone paths are not an indispensable condition for successful recruitment but rather a means to increase the success of recruiters in persuading their nestmates to forage food at a particular location. Our results are relevant to a speciation theory in scent path-laying meliponine bees, such as Scaptotrigona. In addition, the finding that pheromone path-laying bees are able to recruit to food locations even across barriers such as large bodies of water affects tropical pollination ecology and theories on the evolution of resource communication in insect societies with a flying worker caste.
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Abejas/fisiología , Conducta Animal/fisiología , Feromonas/fisiología , Comunicación Animal , Animales , Conducta Alimentaria/fisiología , Vuelo Animal/fisiología , Clima TropicalRESUMEN
Females are often thought to use several cues and more than one modality in selection of a mate, possibly because they offer complementary information on a mate's suitability. In the red mason bee, Osmia rufa, we investigated the criteria a female uses to choose a mating partner. We hypothesized that the female uses male thorax vibrations and size as signs of male viability and male odor for kin discrimination and assessment of genetic relatedness. We therefore compared males that had been accepted by a female for copulation with those rejected, in terms of their size, their immediate precopulatory vibrations (using laser vibrometry), the genetic relatedness of unmated and mated pairs (using microsatellite markers) and emitted volatiles (using chemical analyses). Females showed a preference for intermediate-sized males that were slightly larger than the modal male size. Furthermore, male precopulatory vibration burst duration was significantly longer in males accepted for copulation compared with rejected males. Vibrations may indicate vigor and assure that males selected by females are metabolically active and healthy. Females preferentially copulated with males that were genetically more closely related, possibly to avoid outbreeding depression. Volatiles of the cuticular surface differed significantly between accepted and rejected males in the relative amounts of certain hydrocarbons, although the relationship between male odor and female preference was complex. Females may therefore also use differences in odor bouquet to select among males. Our investigations show that O. rufa females appear to use multiple cues in selecting a male. Future investigations are needed to demonstrate whether odor plays a role in kin recognition and how the multiple cues are integrated in mate choice by females.
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Abejas/fisiología , Conducta de Elección/fisiología , Preferencia en el Apareamiento Animal/fisiología , Animales , Abejas/genética , Tamaño Corporal , Cromatografía de Gases , Femenino , Integumento Común , Masculino , Odorantes/análisis , Filogenia , Vibración , VolatilizaciónRESUMEN
The metatarsal lyriform organ of the spider Cupiennius salei is a vibration detector consisting of 21 cuticular slits supplied by two sensory cells each, one ending in the outer and the other at the inner slit membrane. In search of functional differences between the two cell types due to differences in stimulus transmission, we analyzed (1) the adaptation of responses to electrical stimulation, (2) the thresholds for mechanical stimulation and (3) the representation of male courtship vibrations using intracellular recording and staining techniques. Single- and multi-spiking receptor neurons were found among both cell types, which showed high-pass filter characteristics. Below 100-Hz threshold, tarsal deflections were between 1 degrees and 10 degrees. At higher frequencies, they decreased down to values as small as 0.05 degrees, corresponding to 4.5-nm tarsal deflection in the most sensitive cases. Different slits in the organ and receptor cells with slow or fast adaptation did not differ in this regard. When stimulated with male courtship vibrations, both types of receptor cells again did not differ significantly regarding number of action potentials, latency and synchronization coefficients. Surprisingly, the differences in dendrite coupling were not reflected by the physiological responses of the two cell types innervating the slits.
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Mecanorreceptores/fisiología , Órganos de los Sentidos/citología , Células Receptoras Sensoriales/clasificación , Células Receptoras Sensoriales/fisiología , Arañas/anatomía & histología , Análisis de Varianza , Animales , Biofisica , Biotina/análogos & derivados , Biotina/metabolismo , Cortejo , Femenino , Masculino , Potenciales de la Membrana/fisiología , Técnicas de Placa-Clamp/métodos , Estimulación Física/métodos , Umbral Sensorial/fisiología , VibraciónRESUMEN
Arachnid slit sensilla respond to minute strains in the exoskeleton. After having applied finite element (FE) analysis to simplified arrays of five straight slits (Hössl et al. J Comp Physiol A 193:445-459, 2007) we now present a computational study of the effects of more subtle natural variations in geometry, number and arrangement of slits on the slit face deformations. Our simulations show that even minor variations in these parameters can substantially influence a slit's directional response. Using white-light interferometric measurements of the surface deformations of a lyriform organ, it is shown that planar FE models are capable of predicting the principal characteristics of the mechanical responses. The magnitudes of the measured and calculated slit face deformations are in good agreement. At threshold, they measure between 1.7 and 43 nm. In a lyriform organ and a closely positioned loose group of slits, the detectable range of loads increases to approximately 3.5 times the range of the lyriform organ alone. Stress concentration factors (up to ca. 29) found in the vicinity of the slits were evaluated from the models. They are mitigated due to local thickening of the exocuticle and the arrangement of the chitinous microfibers that prevents the formation of cracks under physiological loading conditions.
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Análisis de Elementos Finitos , Mecanotransducción Celular , Modelos Biológicos , Órganos de los Sentidos/fisiología , Arañas/fisiología , Animales , Fenómenos Biomecánicos , Simulación por Computador , Extremidades , Femenino , Microscopía de Interferencia , Órganos de los Sentidos/anatomía & histología , Umbral Sensorial , Arañas/anatomía & histología , Estrés Mecánico , Soporte de PesoRESUMEN
Stingless bees of the species Trigona spinipes (Fabricius 1793) use their saliva to lay scent trails communicating the location of profitable food sources. Extracts of the cephalic labial glands of the salivary system (not the mandibular glands, however) contain a large amount (approx. 74%) of octyl octanoate. This ester is also found on the scent-marked substrates at the feeding site. We demonstrate octyl octanoate to be a single compound pheromone which induces full trail following behaviour. The identification of the trail pheromone in this widely distributed bee makes it an ideal organism for studying the mechanism of trail following in a day flying insect.
Asunto(s)
Comunicación Animal , Abejas/fisiología , Feromonas/fisiología , Saliva/fisiología , Animales , Abejas/anatomía & histología , Abejas/metabolismo , Caprilatos/metabolismo , Feromonas/química , Saliva/químicaRESUMEN
Atomic force microscopy (AFM) and surface force spectroscopy were applied in live spiders to their joint pad material located distal of the metatarsal lyriform organs, which are highly sensitive vibration sensors. The surface topography of the material is sufficiently smooth to probe the local nanomechanical properties with nanometre elastic deflections. Nanoscale loads were applied in the proximad direction on the distal joint region simulating the natural stimulus situation. The force curves obtained indicate the presence of a soft, liquid-like epicuticular layer (20-40 nm thick) above the pad material, which has much higher stiffness. The Young modulus of the pad material is close to 15 MPa at low frequencies, but increases rapidly with increasing frequencies approximately above 30 Hz to approximately 70 MPa at 112 Hz. The adhesive forces drop sharply by about 40% in the same frequency range. The strong frequency dependence of the elastic modulus indicates the viscoelastic nature of the pad material, its glass transition temperature being close to room temperature (25 +/- 2 degrees C) and, therefore, to its maximized energy absorption from low-frequency mechanical stimuli. These viscoelastic properties of the cuticular pad are suggested to be at least partly responsible for the high-pass characteristics of the vibration sensor's physiological properties demonstrated earlier.