[ULTRASTRUCTURAL ORGANIZATION OF CELLULAR ELEMENTS AND INTERCELLULAR CONNECTIONS IN STATOCYSTS OF TERRESTRIAL PULMONARY SNAIL H. LUCORUM].

The organ of mollusc equilibrium - statocyst appears to be the analogue of acoustic-vestibular system in vertebrate animals. In terrestrial pulmonary snail the epithelial lining of statocyst cavity is created by two types of the cells - a small amount of large cells, provided with kinocilia of sensitive cells and considerably a large number of small supporting or inserted cells, covered with the microvilli. By means of transmission and scanning electron microscopy the ultrastructure and intercellular connections of these cells were studied. The sensitive cells have in a certain way structured cytoplasm, which consists of three layers: ectoplasm, granular layer and hyaloplasm. Myelin-like bodies having the granular, vesicular and drop-like formations in the centre appear to be the special structure of the cytoplasm. In the cytoplasm there are areas, saturated with electron dense glycogen granules. On the electronograms sometimes it is observed how the pinocytic vesicles in the supporting cells are created from the diverticulum of plasmatic membrane of sensitive cells. The boundary areas of plasmatic membrane of adjacent cells (sensitive cells with supporting or supporting cells with the support) are also characterized by the presence of specialized contacts, which are analogous to desmosomes in the epithelial tissues, as well as by the existence of cellular desmosomes, interdigitations. Numerous lacunas have been revealed in the intercellular space, which are connected by the thin tubules and ducts resulting in the formation of a complicated configuration of extensive system of communicating with each other lacunae, which have the exit in statocyst cavity.

[LIGHT AND ELECTRON MICROSCOPY OF CELLULAR ELEMENTS IN STATOCYSTS OF TERRESTRIAL PULMONARY SNAIL HELIX LUCORUM].

Statocysts H. Lucorum appear to be the paired formations having a spherical form and locating on dorsolateral surface of pedal ganglia of subesophageal ganglionic complex. Using the method of reconstruction, as well as the method of scanning electron microscopy on paraffin and semithin slices it has been revealed that the epithelial lining of H. Lucorum appears to be a spatially ordered complex, consisting of 13 cell assemblies. in each of them one sensory cell is surrounded by the satellite cells. The sensory cell has a star shape on the front pole of statocyst because of many protoplasmic processes branching off from its body. The other 12 cells have polygonal shape and form three belts around the inside perimeter of statocyst: interior, middle or equatorial and posterior. There are 4 cells in each belt. The sensory cells forming anterior and posterior belts turned out to be displaced in relation to the cells of equatorial belt and for this reason the whole cell structure resembles a brickwork. The distribution of sensory cells on the inner surfce of statocyst creates the structural polarization of this formtion. First of all, this is evidenced by the presence of the sensory cell on the anterior pole of statocyst and by the absence of those on opposite, posterior pole. The topography of each sensory cell on the inner surface of statocyst, the sizes and shapes of the nuclei, the quantity of nucleoli, the peculiarities of cytoplasm vacuolation and interrelation of sensory and supporting cells have been established.

[Light microscopy of statocyst cell elements from Helix lucorum (space experiment aboard the orbital station "MIR")].

Statocyst epithelial lining of terrestrial pulmonary snail Helix lucorum is a spatially arranged structure consisting of 13 cell ensembles. Each ensemble has a sensory cell surrounded by companion cells. The sensory cell on the anterior statocyst pole is star-shaped due to multiple protoplasmatic protrusions on its body. The remaining 12 polygon-shaped cells form 3 tires along the statocyst internal perimeter: anterior, middle or equatorial and posterior. There are 4 cells in each tire. Topography of every sensory cell on the statocyst internal surface was described as well as cell nuclei size and form, nucleoli number and patterns of cytoplasm vacuolization. Space free of sensory cells is occupied by supporting or intercalary cells. Exposure to space microgravity over 40, 43, 102 and 135 days aboard the orbital station MIR affected morphology of the sensory cells. Specifically, this appeared as reductions in cell height and, consequently, extension of the statocyst cavity internal diameter and volume in the space-flown snails.

[Biological experiments in microgravity: equilibrium function].

The review deals with the investigations of structural and functional modifications in the equilibrium organ (EO) in invertebrates (coelenterates, shells, crustaceans and insects) and vertebrates (fishes, amphibians, rats, primates) on different ontogenetic stages in the condition of microgravity and during readaptation to the Earth's gravity. Results of the investigations detail the adaptive strategy of terrestrial organism in the environment lacking the gravitational components that leads to the discrepancy of an inner model of the body-environment schema constructed by the central nervous system at 1 g and the novel reality. It is manifested by ataxic behavior and increased graviceptors' afferentation against efferent system inactivation. The new condition is defined as a sensibilization phase ensued by the eluding phase: behavior obeys the innate motion strategy, whereas graviceptors' afferentation decreases due to activation of the efferent system. Readaptation to 1 G takes several to 50 days and proceeds as a sequence of slow in motion behavior, ataxia and vestibular sensitization. Reactivity of the gravitosensory system to microgravity was found to be age-dependent. Gain in the EO inertial mass in microgravity and reduction with return to 1 g indicates gravity relevance to EO genesis.

[Plasticity of stastocyst inertial mass in terraneous gastropods helix lucorum and pomatias rivulare in altering gravitational field (microgravity, hypergravity)].

Light and scanning electron microscopy was used to study the morphological parameters and ultrastructure of Helix lucorum statocysts and statoliths in Pomatias rivulare statocysts after 56, 93 and 110-day exposure to microgravity aboard the ISS. Increased gravity was simulated by 30-d centrifugation at 6 g. On the first day of recovery, many statoconia and statoliths were found to carry numerous warts. Moreover, statoconia grew in number significantly as compared with the ground control. On the contrary centrifugation caused massive destruction of statoconia. In a month after orbital flight and centrifugation morphology of both statoconia and stastoliths was nearly normal. These results evidence, that the gravitational field is an important factor for the abiotic medium responsible for building up the inertial mass in the equilibrium organ of animals.

[Equipment for biological experiments with snails aboard piloted orbital stations].

To fly biological experiments aboard piloted orbital stations, research equipment was built up of an incubation container, filter system and automatic temperature controller. Investigations included analysis of the makeup and concentrations of gases produced by animals (snails) during biocycle, and emitted after death. Filters are chemisorption active fibrous materials (AFM) with high sorption rate and water receptivity (cation exchange fiber VION-KN-1 and anion exchange fiber VION-AS-1), and water-repellent carbon adsorbent SKLTS. AFM filters were effective in air cleaning and practically excluded ingress of chemical substances from the container into cabin atmosphere over more than 100 days.

Morphological peculiarities statoconia in statocysts of terrestrial pulmonary snail Helix lucorum.

Vast majority of statoconia in statocysts of Helix lucorum are of oval shape and have smooth surface. Each statoconium in its central part has a nucleus, a spherical mass of 1.5 µ in diameter, surrounded by concentric structures. Minority of statoconia are of subcircular, elongated, rectangular, triangular, irregular, and sometimes fanciful shape and are structured around several nuclei or around small statoconia consolidated by shared growth layers. Apart from statoconia, spherical formations lacking mineral composition of 0.3-2.5 µ in diameter were also found in the statocyst cavity. Similar formations were found in vacuoles of sensory cells of statocyst epithelial lining. It is hypothesized that statoconium nuclei and growth layers around them are of different origin: the nuclei are formed by statocyst sensory cells, while the mineral component is a result of activity of supporting cells.

[The electrovestibular reactions]. / Elektrovestibuliarnye reaktsii.

The article reviews results of studies of electric stimulation of the labyrinth in humans and animals. Electrostimulation of this organ either amplifies or inhibits vestibular reactions till complete suppression of both labyrinths revealed by muteness of labyrinth reflexes and loss of the ability to swim. Uniateral labyrinth destruction can be remedied by cathodal stimulation. Results of the author's investigations implicate current effectivness in controlling the vestibular input including cases of severe vestibular disfunction, e.g. death of labyrinth receptor formations, bilateral tumors, mechanic damage of nerve 8 and others.

Electrical stimulation of labyrinths and vestibular reactions.

Stimulation of the labyrinth in rabbits and guinea pigs with ascending (cathode) or descending direct current (anode) induced deviation and nystagmus of the eyes, head turn and tilt, and other reactions. Anodization of both labyrinths was followed by the disappearance of reflexes, suppression of vestibular reactions to adequate stimulation, and loss of swimming ability. The state of animals returned to normal after breaking the electric circuit.

[The effect of microgravity on the state of compensation in one-side labyrinthisized newts]. / Vliianie nevesomosti na sostoianie kompensatsii u odnostoronne delabirintirovannykh tritonov.

Experiments were performed on newts Pleurodeles waltii subjected to one-side labyrinthectomy 1, 2, 5, and 6 mos prior to a spaceflight on board the orbital complex Mir. Duration of the exposure in microgravity was 8, 31 and 73 days. Destruction of one labyrinth in the newts brought about a number of characteristic reactions, i.e. rollover, circling and sideway movements with the head on a solid substrate and spinning about the longitudinal body axis when in water. These reactions were directed exclusively towards the labyrinthectomized side, their intensity faded, and typically in 1.5 mos. following surgery the motor activity of the newts was close to the norm. In the weight-free conditions the animals appeared to lose compensation they had formed on Earth. During 1.5 days of return the newts were unable to walk and their heads were turned towards the labyrinthectomized side. Immersed in water, the animals started spinning also towards this side. Two days later, they regained the ability to walk without assistance but with evident signs of ataxia. In 40 days after flight, the animals still exhibited locomotor disturbances during walk and swimming.

[Electrophysiological characteristics of the statocyst receptor cells in snails Helix lucorum following exposure to microgravity in unmanned spacecraft "Foton" and piloted space station "Mir"]. / Elektrofiziologicheskie kharakteristiki retseptornykh kletok statotsista ulitok Helix lucorum, éksponirovannykh v nevesomosti na avtomaticheskom kosmicheskom apparate "Foton" i pilotiruemom orbital'nom komplekse "Mir".

Following 16-, 53-, 73- and 163-d orbiting, the background pulsation of the statocyst receptor cells (statoreceptors), the equilibrium organ in land pulmonary snails Helix lucorum, was found increased. Alteration of the body spatial position (2-min tilt at 90 degrees toward the upsilateral side) in the space-flown snails was noted to occur with delayed dying away of the pulsation rate. During stepwise (30 degrees) tilts in the range from 0 to 180 degrees, the maximum of the reaction was shifted by 30 degrees from the gravitational vertical; during dynamic tilting (sine decaying oscillations) the reaction subsides more rapidly as compared with the control animals. Absence of changes in pulsation of receptor cells in statocysts deprived of feedback with the nervous centers may be suggestive of inactivation of the efferent innervation in consequence of exposure to microgravity.

Effect of static and dynamic influences on receptors of equilibrium organ of Helix lucorum after 163-day orbital flight in "Mir" station.

A 163-day orbital flight increased the baseline impulse activity of statocyst receptor cells in terrestrial pulmonata snail Helix lucorum. The maximum of reaction to step static stimuli (changes in body position in the range from 0 to 180 degrees) was significantly shifted by 30 degrees from gravitational vertical, while the reaction to dynamic stimuli (dumped sinusoidal oscillations) faded more rapidly than in control animals.