BackgroundThe mollusk statocyst is a mechanosensing organ detecting the animal's orientation with respect to gravity. This system has clear similarities to its vertebrate counterparts: a weight-lending mass, an epithelial layer containing small supporting cells and the large sensory hair cells, and an output eliciting compensatory body reflexes to perturbations.Methodology/Principal FindingsIn terrestrial gastropod snail we studied the impact of 16- (Foton M-2) and 12-day (Foton M-3) exposure to microgravity in unmanned orbital missions on: (i) the whole animal behavior (Helix lucorum L.), (ii) the statoreceptor responses to tilt in an isolated neural preparation (Helix lucorum L.), and (iii) the differential expression of the Helix pedal peptide (HPep) and the tetrapeptide FMRFamide genes in neural structures (Helix aspersa L.). Experiments were performed 13–42 hours after return to Earth. Latency of body re-orientation to sudden 90° head-down pitch was significantly reduced in postflight snails indicating an enhanced negative gravitaxis response. Statoreceptor responses to tilt in postflight snails were independent of motion direction, in contrast to a directional preference observed in control animals. Positive relation between tilt velocity and firing rate was observed in both control and postflight snails, but the response magnitude was significantly larger in postflight snails indicating an enhanced sensitivity to acceleration. A significant increase in mRNA expression of the gene encoding HPep, a peptide linked to ciliary beating, in statoreceptors was observed in postflight snails; no differential expression of the gene encoding FMRFamide, a possible neurotransmission modulator, was observed.Conclusions/SignificanceUpregulation of statocyst function in snails following microgravity exposure parallels that observed in vertebrates suggesting fundamental principles underlie gravi-sensing and the organism's ability to adapt to gravity changes. This simple animal model offers the possibility to describe general subcellular mechanisms of nervous system's response to conditions on Earth and in space.
The utricle provides the vestibular reflex pathways with the sensory codes of inertial acceleration of self-motion and head orientation with respect to gravity to control balance and equilibrium. Here we present an anatomical description of this structure in the adult oyster toadfish and establish a morphological basis for interpretation of subsequent functional studies. Light, scanning, and transmission electron microscopy techniques were applied to visualize the sensory epithelium at varying levels of detail, its neural innervation and its synaptic organization. Scanning electron microscopy was used to visualize otolith mass and morphological polarization patterns of hair cells. Afferent nerve fibers were visualized following labeling with biocytin, and light microscope images were used to make three-dimensional (3-D) reconstructions of individual labeled afferents to identify dendritic morphology with respect to epithelial location. Transmission electron micrographs were compiled to create a serial 3-D reconstruction of a labeled afferent over a segment of its dendritic field and to examine the cell-afferent synaptic contacts. Major observations are: a well-defined striola, medial and lateral extra-striolar regions with a zonal organization of hair bundles; prominent lacinia projecting laterally; dependence of hair cell density on macular location; narrow afferent dendritic fields that follow the hair bundle polarization; synaptic specializations issued by afferents are typically directed towards a limited number of 7-13 hair cells, but larger dendritic fields in the medial extra-striola can be associated with > 20 hair cells also; and hair cell synaptic bodies can be confined to only an individual afferent or can synapse upon several afferents.
Gravity plays a key role in shaping the vestibular sensitivity (VS) of terrestrial organisms. We studied VS changes in the statocyst of the gastropod Helix aspersa immediately after 4-, 16-, and 32-day exposures to a 1.4G hypergravic field or following a 7-day recovery period. In the same animals we measured latencies of behavioral "negative gravitaxis" responses to a head-down pitch before and after centrifugation and found significant delays after 16- and 32-day runs. In an isolated neural preparation we recorded the electrophysiological responses of the statocyst nerve to static tilt (±19°) and sinusoids (±12°; 0.1 Hz). Spike sorting software was used to separate individual sensory cells' patterns out of a common trace. In correspondence with behavior we observed a VS decrease in animals after 16- (p < 0.05) and 32-day (p < 0.01) centrifugations. These findings reveal the capability of statoreceptors to adjust their sensitivity in response to a prolonged change in the force of gravity. Interestingly, background discharge rate increased after 16 and 32 days in hypergravity and continued to rise through the recovery period. This result indicates that adaptive mechanisms to novel gravity levels were long lasting, and re-adaptation from hypergravity is a more complex process than just "return to normal".
The vestibular system receives a permanent influence from gravity and reflexively controls equilibrium. If we assume gravity has remained constant during the species' evolution, will its sensory system adapt to abrupt loss of that force? We address this question in the land snail Helix lucorum exposed to 30 days of near weightlessness aboard the Bion-M1 satellite, and studied geotactic behavior of postflight snails, differential gene expressions in statocyst transcriptome, and electrophysiological responses of mechanoreceptors to applied tilts. Each approach revealed plastic changes in the snail's vestibular system assumed in response to spaceflight. Absence of light during the mission also affected statocyst physiology, as revealed by comparison to dark-conditioned control groups. Readaptation to normal tilt responses occurred at ~20 h following return to Earth. Despite the permanence of gravity, the snail responded in a compensatory manner to its loss and readapted once gravity was restored.
The paper presents an algorithm to calculate feasible steady-state conditions of an electrical network. A distinctive feature of the calculation is the consideration of constraints on the temperature of bare conductor and the clearance of a controlled span of overhead line. This consideration is based on forming and solving an equation of conductor heat balance to calculate the conductor temperature depending on the current flowing through the conductor and environmental factors, including the actual values of air pressure and air temperature. The paper presents the results of numerical calculations performed for individual overhead lines and for a test 16-node scheme of electrical network. The results illustrate the algorithm performance and the importance of considering the above constraints.
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