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Circadian rhythms in a long-term duration space flight.

In order to maintain cosmonaut health and performance, it is important for the work-rest schedule to follow human circadian rhythms (CR). What happens with CR in space flight? Investigations of CR in mammals revealed, that the circadian phase in flight is less stable, probably due to a displacement of the range of entrainment, resulting from internal period change (the latter was confirmed on insects). The circadian period may be a gravity-dependent parameter. If so, the basic biological requirement for the day length might be different in weightlessness. On this basis, a higher risk of desynchronosis is expected in a long-duration space flight. As a countermeasure, a non-24-hr day length could be suggested, being close to the internal circadian period (in humans about 25 hr). Taking into account a possible displacement of period in weightlessness, it seems reasonable to establish a flexible work-rest schedule, capable to follow the body temperature CR by means of biofeedback.

Animals↗

Gravimorphogenesis and ultrastructure of the fungus Flammulina velutipes grown in space, on clinostats and under hyper-g conditions.

The D-2-mission provided the facilities to cultivate the higher basidomycete Flammulina velutipes (Agaricales) in space for about 8 days. Gravimorphogenesis of developing fruiting body primordia in weightlessness was documented in comparison to cultures incubated on a 1xg reference centrifuge in space. Chemical fixation of fruiting bodies took place for later ultrastructural analysis. The microgravity grown fruiting bodies exhibited random orientation compared to the 1xg-cultures where fruiting bodies showed exactly negative gravitropic orientation. Weightlessness did not impair fruiting body morphogenesis and growth although flat and helically twisted stipes were observed. Ultrastructural analyses of microgravity-, 1xg- and 20xg-samples did not reveal sedimentable cell components. Gravitropic bending involves growth inhibition at the upper side of a horizontally oriented transition zone, the graviperceptive region of the stipe. The fastest ultrastructural response to the altered direction of the accelerational force is the accumulation of cytosolic vesicles at the lower part of this region. They contribute to the expansion of the central vacuole and therefore to the differential enlargement of the lower side of the stipe.

Basidiomycota↗

Interaction between radiation effects, gravity and other environmental factors in Tribolium confusum.

Multicellular organisms possess homeostatic control systems, which, in responding to changes of the external environment, modify the internal milieu of the organism accordingly, in order to make survival and normal physiological processes possible. This group has studied effects on physiological processes of single and combined environmental factors in the flour beetle, Tribolium confusum. Studies included low- and high-LET radiation, gravity compensation, near-weightlessness in space flight, ambient temperature, atmospheric composition, and magnetic field effects on the growth and development of Tribolium. For somatic effects, there appears to be a "normal physiological range" for each of these environmental variables; moreover, their effects seem independent of each other. When one of the listed environmental factors is near the limits of normal physiological tolerance, however, marked synergism has been observed between the effects of this factor and other environmental stresses. Fertilized Tribolium eggs showed a differential radiosensitivity to external irradiation, and a linear, dose-effect relationship was obtained, as accelerated heavy nitrogen and oxygen ions were used. Synergism was found when Tribolium were irradiated with ionizing radiation and subsequently exposed to temperature either below or above their normal physiological range. Gravity compensation or near-weightlessness in space flight have adverse effects on the development of Tribolium when an ambient temperature is near the higher or lower limit of tolerance. Similar results were observed when pupae were exposed to a combination of oxygen, temperature and magnetic field stresses.

Animals↗

A model for vestibular function in altered gravitational states.

During evolution, the vestibular organ was made to serve mainly two purposes: 1) to guide eye movements during sharp turns, so that the point of fixation in the visual field can be kept steady, a function accomplished by the semicircular canal system and 2) to indicate the terrestrial vertical, so that upright posture and gait can be maintained even in the dark. The otolith system serves the latter purpose. Since the function of the semicircular canal system does not depend on gravity, it is not grossly disturbed by gravitational levels different from 1 g. The proper function of the otolith system depends entirely on the presence of a gravitational force vector of 9.8/m/sec2 directed towards the center of the earth. This system therefore malfunctions when the amplitude of the combined gravito-inertial load is different from 1 g and also when the direction of the sensed gravitational pull is "contaminated" by additional inertial reactive forces as during horizontal acceleration. The effect of such inertial stimulations is probably even stronger in a weightless environment, in which case the background stimulation of terrestrial gravity is absent. Moreover, minor mass differences between the otolithic membranes of the left and right inner ear, even if well compensated on the ground, might lead to malcompensation in weightlessness as well as in hypergravity. A hypothetical model is developed to describe in the central nervous system compensating mechanisms in hypo- and hypergravitational states. The "space-sled" is introduced as a new research tool and recommendations are made for a prophylactic training regimen to reduce or prevent space sickness.

Adaptation, Physiological↗

Is gravity a morphological determinant in plants at the cellular level?

It is not yet known whether higher plants, which have evolved in a gravity oriented environment, can reproduce normally or develop via single cells (somatic or zygotic) and pass normally through all phases of development, tissue differentiation and reproduction through consecutive generations at near-zero g. Most published work deals with directional gravitational stimuli (geotropisms) as they operate on the subsequent growth of preformed plant organs that have acquired the ability to perceive and respond to gravity during development at 1 g. But it should be established whether plant development can proceed normally in the weightless state, especially that critical stage in which single cells produce multicellular units leading to embryos with the growing regions of shoot and root which, in turn, give rise to all the tissues of the plant body. We consider here an experiment that tested whether carrot embryos, capable of developing from cultured somatic cells, could do so under conditions of near-weightlessness. Space flight seemed to have no adverse affects on the induction of such embryos or on the initiation of their organs. But there are still reservations which are discussed. Other systems adaptable to the study of embryogenesis and development in space and to resolve the current reservations are described and their place in future space research is indicated.

Arabidopsis↗

Kinematic and dynamic processes for the control of pointing movements in humans revealed by short-term exposure to microgravity.

The generation of accurate motor commands requires implicit knowledge of both limb and environmental dynamics. The action of gravity on moving limb segments must be taken into account within the motor command, and may affect the limb trajectory chosen to accomplish a given motor task. Exactly how the CNS deals with these gravitoinertial forces remains an open question. Does the CNS measure gravitational forces directly, or are they accommodated in the motor plan by way of internal models of physical laws? In this study five male subjects participated. We measured kinematic and dynamic parameters of upward and downward arm movements executed at two different speeds, in both normal Earth gravity and in the weightless conditions of parabolic flight. Exposure to microgravity affected velocity profiles for both directions and speeds. The shape of velocity profiles (the ratio of maximum to mean velocity) and movement duration both showed transient perturbations initially in microgravity, but returned to normal gravity values with practice in 0 x g. Differences in relative time to peak velocity between upward versus downward movements, persisted for all trial performed in weightlessness. These differences in kinematic profiles and in the torque profiles used to produce them, diminished, however, with practice in 0 x g. These findings lead to the conclusion that the CNS explicitly represents gravitational and inertial forces in the internal models used to generate and execute arm movements. Furthermore, the results suggest that the CNS adapts motor plans to novel environments on different time scales; dynamics adapt first to reproduce standard kinematics, and then kinematics patterns are adapted to optimize dynamics.

Adult↗

Recording heart rate and blood pressure in rats during parabolic flight.

The way in which the cardiovascular system adapts to weightlessness is still under discussion. No data are yet available on the responses of rats during space flight, although this animal is commonly used in simulation studies. We have designed and tested a protocol to study the short term responses of the cardiovascular system to weightlessness during parabolic flight. A telemetry system was used to measure heart rate (HR) and blood pressure. It was possible to collect and record radio-signals without any interference. Microgravity caused a reduction in HR, an increase in mean arterial pressure (MAP, 7%), and a non-significant decrease in central venous pressure (CVP, 13%). The change in CVP was similar to the decrease observed in human space flight. This type of study may also be feasible for longer exposure of rats to microgravity (space flight).

Animals↗

Does gravity play an essential role in the asymmetrical visual perception of vertical and horizontal line length?

The eye perceives the length of vertical and horizontal lines with an inherent asymmetry. A vertical line having the same length as a horizontal one is usually perceived to be longer. In this experimental investigation we tested the hypothesis that gravity has a direct role in producing the observed perceptual asymmetry. To this end we performed experiments in weightlessness during long-orbital space flights onboard the MIR station. Subjects performed a psychophysical task in which the length of a visually-presented vertical line was adjusted to match the length of a horizontal reference. On Earth, almost all subjects produce errors in adjusting the length of the vertical line, consistently under-estimating the length of the horizontal reference. The asymmetry of perception of the line lengths persisted in weightlessness. From these results we conclude that the phenomena of asymmetry of perception of the lengths of vertical and horizontal lines is not dependent on gravity, but is instead defined by properties of the system of internal representation. Grant numbers: 99-04-48450.

Adaptation, Physiological↗

ELITE-S2: the multifactorial movement analysis facility for the International Space Station.

Experimental observations of adaptation processes of the motor control system to altered gravity conditions can provide useful elements to the investigations on the mechanisms underlying motor control of human subject. The microgravity environment obtained on orbital flights represents a unique experimental condition for the monitoring of motor adaptation. The research in motor control exploits the changes caused by microgravity on the overall sensorimotor process, due to the impairment of the sensory systems whose function depends upon the presence of the gravity vector. Motor control in microgravity has been investigated during parabolic flights and short-term space missions, in particular for analysis of movement-posture co-ordination when equilibrium is no longer a constraint. Analysis of long-term adaptation would also be very interesting, calling for long-term body motion observations during the process of complete motor adaptation to the weightlessness environment. ELITE-S2 is an innovative facility for quantitative human movement analysis in weightless conditions onboard the International Space Station (ISS). ELITE-S2 is being developed by the Italian Space Agency, ASI is to be delivering the flight models to NASA to be included in an expressed rack in US Lab Module in February 2004. First mission is currently planned for summer 2004 (increment 10 ULF 2 ISS).

Adaptation, Physiological↗

Neurobiological responses of fish to altered gravity conditions: a review.

In vertebrates (including man), altered gravitational environments such as weightlessness can induce malfunctions of the inner ears, based on an irregular dislocation of the inner ear otoliths from the corresponding sensory epithelia. This dislocation leads to an illusionary tilt, since the otolithic inputs are not confirmed by the other sensory organs, which results in an intersensory conflict. Vertebrates in the orbit therefore face severe orientation problems. In humans, the intersensory conflict may additionally lead to a malaise, commonly referred to as space motion sickness (SMS). During the first days at weightlessness, the orientation problems (and SMS) disappear, since the brain develops a new compensatory interpretation of the available sensory data. The present review reports on the neurobiological responses--particularly of fish--observed at altered gravitational states, concerning behaviour and neuroplastic reactivities.

Animals↗

Arm end-point trajectories under normal and micro-gravity environments.

The purpose of the present experiment was to study the way in which the CNS represents gravitational force during vertical arm pointing movements. Movements in upward and downward directions were executed by two cosmonauts in normal-gravity and weightlessness. Analyses focused upon finger kinematics in the sagittal plane. In normal-gravity, downward direction movements showed smaller curvatures and greater relative times to peak velocity (AT/MT) when compared with upward direction movements. Data from the weightlessness experiments showed that whilst downward movements decreased their curvature during space flight, curvatures of upward movements changed slightly. Furthermore, AT/MT was modified during the first days in micro-gravity for both directions, recovering, however, to pre-flight values after 18 days in space. Results from the present study, provide evidence that gravitational force is centrally treated constituting an important component of the motor plan for vertical arm movements.

Aerospace Medicine↗

DARA vestibular equipment onboard MIR.

In space, the weightless environment provides a different stimulus to the otolith organs of the vestibular system, and the resulting signals no longer correspond with the visual and other sensory signals sent to the brain. This signal conflict causes disorientation. To study this and also to understand the vestibular adaptation to weightlessness, DARA has developed scientific equipment for vestibular and visuo-oculomotoric investigations. Especially, two video-oculography systems (monocular--VOG--and binocular--BIVOG, respectively) as well as stimuli such as an optokinetic stimulation device have successfully been employed onboard MIR in the frame of national and European missions since 1992. The monocular VOG was used by Klaus Flade during the MIR '92 mission, by Victor Polyakov during his record 15 months stay onboard MIR in 1993/94 as well as by Ulf Merbold during EUROMIR '94. The binocular version was used by Thomas Reiter and Sergej Avdeyev during the 6 months EUROMIR '95 mission. PIs of the various experiments include H. Scherer and A. Clarke (FU Berlin), M. Dieterichs and S. Krafczyk (LMU Munchen) from Germany as well as C.H. Markham and S.G. Diamond from the United States. Video-Oculography (VOG) is a technique for examining the function of the human balance system located in the inner ear (vestibular system) and the visio-oculomotor interactions of the vestibular organ. The human eye movements are measured, recorded and evaluated by state-of-the-art video techniques. The method was first conceived and designed at the Vestibular Research Laboratory of the ENT Clinic in Steglitz, FU Berlin (A. Clarke, H. Scherer). Kayser-Threde developed, manufactured and tested the facilities for space application under contract to DARA. Evaluation software was first provided by the ENT Clinic, Berlin, later by our subcontractor Sensomotoric Instruments (SMI), Teltow. Optokinetic hardware to support visuo-oculomotoric investigations, has been shipped to MIR for EUROMIR '95 and has successfully been used in conjunction with VOG by ESA astronaut Thomas Reiter. Most recently, BIVOG aboard MIR will be reused in the frame of German/Russian joint experiment sessions employing two Russian cosmonauts from August 1997 to January 1998.

Aerospace Medicine↗

Unexpected renal responses in space.

Urine output in astronauts following ingestion of an oral water load was low in space on the Russian space station Mir and less than during simulation by 6 degrees head-down bed rest. This surprising observation shows that the effects of gravity and weightlessness on fluid volume regulation are not well understood and that the head-down bed-rest model does not simulate the effects of weightlessness on renal water handling.

Adult↗

Neurobiology of fish under altered gravity conditions.

In vertebrates (including man), an altered gravitational environment such as weightlessness can induce malfunction of the inner ear, based on an irregular dislocation of the otoliths from the corresponding sensory epithelia. This dislocation leads to an illusionary tilt, since the otolithic inputs are not in register with other sensory organs. This results in an intersensory conflict. Vertebrates in orbit therefore face severe orientation problems. In humans, the intersensory conflict may additionally lead to a malaise, commonly referred to as space motion sickness (SMS). During the first days in weightlessness, the orientation problems (and SMS) disappear, since the brain develops a new compensatory interpretation of the available sensory data. The present review reports the neurobiological responses-particularly in fish-observed at altered gravitational states, concerning behaviour and neuroplastic reactivities. Recent investigations employing microgravity (spaceflight, parabolic aircraft flights, clinostat) and hyper-gravity (laboratory centrifuges as ground based research tools) yielded clues and insights into the understanding of the respective basic phenomena. The possible sources of human space sickness (a kinetosis) and of the space adaptation syndrome (when a sensory reinterpretation of gravitational and visual cues takes place) are particularly highlighted with regard to the functional significance of bilaterally asymmetric otoliths (weight, size).

Animals↗

The sensorimotor and cognitive integration of gravity.

In order to demonstrate that gravity is not only a load acting locally and continuously on the body limbs, but is also used by higher levels of the nervous system as a dynamic orienting reference for the elaboration of the motor act, a review of several experiments conducted both in 1 g and 0 g are presented. During various locomotor tasks, the strategy that consists of stabilizing the head with respect to gravity illustrates one of the solutions used by the CNS to optimize the control of dynamic equilibrium. A question which remains to be solved when considering experimental results obtained in weightlessness concerns, however, the maintenance of motor schema that has evolved under normal gravity. Results have suggested that the concept of conservative processes, that would adapt postural control to weightlessness by using previously learned innate strategies, must be reconsidered during goal-oriented tasks. In fact, it is proposed that when conservative processes and existing solutions derived from a repertoire of terrestrial postural strategies do not provide efficient output, the CNS has to create novel strategies through a slow learning process. As with the study of postural control, three-dimensional arm reaching movements also illustrate the central representation of gravity. Indeed, gravity can be regarded as either initiating or braking arm movements and, consequently, may be represented in the motor command at the planning level. Finally, from a prospective point of view, there is a need to determine new experimental paradigms in order to study the specific motor control of man in space. It is suggested that the formulation of experimental paradigms should not consider man in space simply as a terrestrial biped.

Arm↗

The effect of gravity on human recognition of disoriented objects.

The role of non-visual cues, and particularly of signals tied to the direction of gravity, in the mechanisms of recognition of disoriented objects is reviewed. In spite of a limited number of studies, object recognition does not seem dramatically altered by weightlessness and astronauts can adapt to this novel environment. Particularly, mental rotation strategy can still be used in weightlessness with dynamic parameters relatively unchanged. Similarly, spatial coordinate assignment can be performed adequately under different gravitational conditions. However, signals related to gravity direction seem to be integrated in the early stages of visual processing. Thus, performances in symmetry detection tasks and visual search tasks are influenced by the gravito-inertial conditions in which experience are done. Functional roles of such a multisensory convergence on cortical visual neurons, partly confirmed by neurophysiological studies, are proposed.

Form Perception↗

Pleurodeles waltl, amphibian, Urodele, is a suitable biological model for embryological and physiological space experiments on a vertebrate.

Pleurodeles waltl (amphibian, Urodele) is an appropriate biological model for space experiments on a vertebrate. One reason for interest in this animal concerns the study of the effects of absence of gravity on embryonic development. First, after mating (on Earth) the females retain live, functional sperm in their cloacum for up to 5 months, allowing normal in vivo fertilisation after hormonal stimulation. Second, their development is slow, which allows analyses of all the key stages of ontogenesis from the oocyte to swimming tailbud embryos or larvae. We have performed detailed studies and analyses of the effects of weightlessness on amphibian Pleurodeles embryos, fertilised and allowed to develop until the swimming larvae stage. These experiments were performed in space during three missions on the MIR-station: FERTILE I, FERTILE II and NEUROGENESIS respectively in 1996, 1998 and 1999. We show that in microgravity abnormalities appeared at specific stages of development compared to 1g-centrifuge control embryos and 1g-ground control embryos. In this report we describe abnormalities occurring in the central nervous system. These modifications occur during the neurulation process (delay in the closure of the neural tube and failure of closure of this tube in the cephalic area) and at the early tailbud stage (microcephaly observed in 40% of the microgravity-embryos). However, if acephalic and microcephalic embryos are not taken into account, these abnormalities did not disturb further morphological, biochemical and functional development and the embryos were able to regulate and a majority of normal hatching and swimming larvae were obtained in weightlessness with a developmental time-course equivalent to that of 1g-centrifuge control embryos (on the MIR station) and 1g-ground control embryos.

Animals↗

Gravitational neurobiology of fish.

In vertebrates (including man), altered gravitational environments such as weightlessness can induce malfunctions of the inner ears, based on irregular movements of the semicircular cristae or on dislocations of the inner ear otoliths from the corresponding sensory epithelia. This will lead to illusionary tilts, since the vestibular inputs are not confirmed by the other sensory organs, which results in an intersensory conflict. Vertebrates in orbit therefore face severe orientation problems. In humans, the intersensory conflict may additionally lead to a malaise, commonly referred to as space motion sickness (SMS), a kinetosis. During the first days at weightlessness, the orientation problems (and SMS) disappear, since the brain develops a new compensatory interpretation of the available sensory data. The present review reports on the neurobiological responses--particularly of fish--observed at altered gravitational states, concerning behaviour and neuroplastic reactivities. Recent investigations employing microgravity (spaceflight, parabolic aircraft flights, clinostat) and hyper-gravity (laboratory centrifuges as ground based research tools) yielded clues and insights into the understanding of the respective basic phenomena.

Animals↗