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Effects of Gravity on Insect Circadian Rhythmicity

Circadian rhythms - endogenous daily rhythmic fluctuations in virtually all characteristics of life - are generated and coordinated by the circadian timing system (CTS). The CTS is synchronized to the external 24-hour day by time cues such as the light/dark cycle. In an environment without time cues, the length of an animal's day is determined by the period of its internal pacemaker (tau) and the animal is said to be free-running. All life on earth evolved under the solar day; the CTS exists as an adaptation that allows organisms to anticipate and to prepare for rhythmic environmental fluctuations. All life on earth also evolved under the force of earth's gravitational environment. While it is therefore not surprising that changes in the lighting environment affect the CTS, it is surprising that changes in the gravitational environment would do so. However, recent data from one of our laboratories using the brn-3.1 knockout mouse revealed that this model, which lacks the sensory receptor hair cells within the neurovestibular system, does not respond to exposure to a hyperdynamic environment in the same fashion as normal mice. The brn-3.1 mice did not show the expected suppression of circadian rhythmicity shown by control mice exposed to 2G. Exposure to altered ambient force environments affects the amplitude, mean and timing of circadian rhythms in species from unicellular organisms to man. In addition, there is a circadian influence on the homeostatic response to acute 2G acceleration and pulses of 2G can act as a time cue, synchronizing the CTS. This is of significance because maintenance of internal and external temporal coordination is critical for normal physiological and psychological function. Typically, during adaptation to an increased gravitational environment (+G), an initial acute reaction is followed by adaptation and, eventually, a new steady state (14-16), which can take weeks to months to establish. Until the development of space stations, exposure to microgravity was, of necessity, relatively short in duration. In early spaceflight experiments an organism's internal rhythms often expressed periods that were different from each other, even in the presence of a 24.0 hour light-dark cycle, suggesting that the organism was experiencing internal desynchronization (17, 18). In (micro)G, the body temperature rhythm was delayed with respect to other body rhythms and to the light-dark cycle in rhesus macaques (19) and man (20, 21). In the absence of a light-dark cycle, the circadian rhythm of spore formation persisted in Neurospora crassa, however, both the variability and average period of the rhythm increased (22). The beetle Trigonoscelis gigas, exhibited changes in period during and following 11-13 days in (micro)G (23, 24). Resynchronization of the urinary calcium rhythm following a 1800 phase shift of the LID cycle was retarded in rats exposed to (micro)G compared to 1G controls (25). With the development of the Russian Mir Space Station, long-term controlled microgravity exposure became possible. We recorded activity rhythms from black-bodied Tenebrionid beetles, Trigonoscelis gigas, in (micro)G (spaceflight). Each insect was housed individually within an activity monitor (26) and data (activity counts) were collected and stored in five-minute bins. Thirty-two individual activity monitors were housed within each of 2 experimental kits. The beetles within each kit were divided into two groups and the lighting was controlled separately for each group.

Hoban-Higgins, Tana M.↗

Microbial Response to Microgravity and Other Low Shear Environments

Microbial existence and survival requires the ability to sense and respond to environmental changes, including changes in physical forces. This is because microbes inhabit an amazingly diverse range of ecological niches and therefore must constantly adapt to a wide variety of changing environmental conditions, including alterations in temperature, pH, nutrient availability, oxygen levels, and osmotic pressure gradients. Microbes sense their environment through a variety of sensors and receptors which serve to integrate the different signals into the appropriate cellular response(s) that is optimal for survival. While numerous environmental stimuli have been examined for their effect on microorganisms, effects due to changes in mechanical and/or physical forces are also becoming increasingly apparent. Recently, several important studies have demonstrated a key role for microgravity and the low fluid shear dynamics associated with microgravity in the regulation of microbial gene expression, physiology and pathogenesis. The mechanosensory response of microorganisms to these environmental signals, which are relevant to those encountered during microbial life cycles on Earth, may provide insight into their adaptations to physiologically relevant conditions and may ultimately lead to eludicidation of the mechanisms important for mechanosensory transduction in living cells. This review summarizes the recent and potential future research trends aimed at understanding the effect of changes in mechanical forces that occur in microgravity and other low shear environments on different microbial parameters. The results of these studies provide an important step towards understanding how microbes integrate information from multiple mechanical stimuli to an appropriate physiological response.

Nickerson, C.↗

Adaptation of primate vestibuloocular reflex to altered peripheral vestibular inputs. I. Frequency-specific recovery of horizontal VOR after inactivation of the lateral semicircular canals

1. The adaptive plasticity of the vestibuloocular reflex (VOR) following a selective lesion of the peripheral vestibular organs was investigated in rhesus monkeys whose lateral semicircular canals were inactivated by plugging of the canal lumen in both ears. Gain and phase of horizontal, vertical, and torsional slow-phase eye velocity were determined from three-dimensional eye movement recordings obtained acutely after the plugging operation, as well as in regular intervals up to 10 mo later. 2. Acutely after plugging, horizontal VOR was minimal during yaw rotation with gains of < 0.1 at all frequencies. Horizontal VOR gain gradually increased over time, reaching gains of 0.4-0.5 for yaw oscillations at 1.1 Hz approximately 5 mo after lateral canal inactivation. This response recovery was strongly frequency dependent: horizontal VOR gains were largest at the highest frequency tested and progressively decreased for lower frequencies. Below approximately 0.1 Hz, no consistent horizontal VOR could be elicited even 10 mo after plugging. 3. The frequency-dependent changes in gain paralleled changes in horizontal VOR phase. Below approximately 0.1-0.05 Hz large phase leads were present, similarly as in semicircular canal primary afferents. Smaller phase leads were also present at higher frequencies, particularly at 1.1 Hz (the highest frequency tested). 4. Consistent with the afferent-like dynamics of the adapted horizontal VOR, per- and postrotatory horizontal responses to constant-velocity yaw rotations were short lasting. Time constants of the slow-phase eye velocity envelope of the horizontal postrotatory nystagmus were approximately 2 s. Nonetheless, a consistent horizontal optokinetic afternystagmus was evoked in plugged animals. 5. A torsional component that was absent in intact animals was consistently present during yaw rotation acutely after lateral canal inactivation and remained approximately constant thereafter. The frequency response characteristics of this torsional component resembled those of the adapted horizontal slow-phase responses: gain decreased and large phase leads were introduced at frequencies below approximately 0.05-0.1 Hz. Torsional responses elicited by roll oscillations in supine position, on the other hand, were indistinguishable in their dynamics from intact animals. No consistent vertical nystagmus was elicited during yaw rotation. 6. Our results show that there is a slow, frequency-specific recovery of horizontal VOR after selective inactivation of the lateral semicircular canals. Both the spatial organization and the dynamic properties of the adapted VOR responses are distinctly different from responses in intact animals, suggesting complex changes in the underlying vestibuloocular circuitry.

NASA Program Space Physiology and Countermeasures↗

Effect of a 30-day isolation stress on calcium, phosphorus and other excretory products in an unrestrained chimpanzee.

An unrestrained chimpanzee was studied in an isolation chamber and in his home cage environment. The study consisted of 49 urine collection days (14 days pre-, 5 days post- and 30 days of isolation), and then of 10 days in the home cage. Dietary intake, urine and fecal data were obtained. The effect of isolation on various excretory parameters was studied. Urine samples were analyzed for volume, osmolarity, creatinine, creatine, urea-N, 17-hydroxy corticosteroids, VMA, calcium and inorganic phosphorus. One way analyses of variance performed on the urinary excretion parameters showed all except creatinine excretion to vary significantly during periods of the study. The changes observed in calcium and phosphorus were highly significant. The data suggests that the calcium to phosphorus excretion ratio might serve as a physiological stress indicator of Selye's adaptation syndrome (period of resistance).

Sabbot, I. M.↗

The internal nature of functional disturbances arising in the body under the effect of extreme factors

Physiological factors in human tolerance and adaptation to pressurized cabin atmospheres during manned space flight are analyzed. It is shown that individuals display different degrees of functional changes and disturbances despite identical indices of compensation by the body of external factors. The significance of reactions at the cellular tissue level to factors of an altered gaseous environment is emphasized.

Gramenitskiy, P. M.↗

Age-related changes in human vestibulo-ocular reflexes: Sinusoidal rotation and caloric tests

The dynamic response properties of horizontal vestibulo-ocular reflex (VOR) were characterized in 216 human subjects ranging in age from 7 to 81 years. The object of this cross-sectional study was to determine the effects of aging on VOR dynamics, and to identify the distributions of parameters which describe VOR responses to caloric and to sinusoidal rotational stimuli in a putatively normal population. Caloric test parameters showed no consistent trend with age. Rotation test parameters showed declining response amplitude and slightly less compensatory response phase with increasing age. The magnitudes of these changes were not large relative to the variability within the population. The age-related trends in VOR were not consistent with the anatomic changes in the periphery reported by others which showed an increasing rate of peripheral hair cell and nerve fiber loss in subjects over 55 years. The poor correlation between physiological and anatomical data suggest that adaptive mechanisms in the central nervous system are important in maintaining the VOR.

Peterka, R. J.↗

Biomedical Polar Research Workshop Minutes

This workshop was conducted to provide a background of NASA and National Science Foundation goals, an overview of previous and current biomedical research, and a discussion about areas of potential future joint activities. The objectives of the joint research were: (1) to develop an understanding of the physiological, psychological, and behavioral alterations and adaptations to extreme environments of the polar regions; (2) to ensure the health, well-being, and performance of humans in these environments; and (3) to promote the application of biomedical research to improve the quality of life in all environments.

Source record↗

Cardiopulmonary discipline science plan

Life sciences research in the cardiopulmonary discipline must identify possible consequences of space flight on the cardiopulmonary system, understand the mechanisms of these effects, and develop effective and operationally practical countermeasures to protect crewmembers inflight and upon return to a gravitational environment. The long-range goal of the NASA Cardiopulmonary Discipline Research Program is to foster research to better understand the acute and long-term cardiovascular and pulmonary adaptation to space and to develop physiological countermeasures to ensure crew health in space and on return to Earth. The purpose of this Discipline Plan is to provide a conceptual strategy for NASA's Life Sciences Division research and development activities in the comprehensive area of cardiopulmonary sciences. It covers the significant research areas critical to NASA's programmatic requirements for the Extended-Duration Orbiter, Space Station Freedom, and exploration mission science activities. These science activities include ground-based and flight; basic, applied, and operational; and animal and human research and development. This document summarizes the current status of the program, outlines available knowledge, establishes goals and objectives, identifies science priorities, and defines critical questions in the subdiscipline areas of both cardiovascular and pulmonary function. It contains a general plan that will be used by both NASA Headquarters Program Offices and the field centers to review and plan basic, applied, and operational (intramural and extramural) research and development activities in this area.

Source record↗

Urinary excretion of LH and testosterone from male rats during exposure to increased gravity: post-spaceflight and centrifugation

A dissociation between plasma luteinizing hormone (LH) and testosterone (T) appears to exist during exposure to altered gravity. The pulsatile nature of LH release and the diurnal variability of T secretion may mask or bias the effects of altered gravity on the pituitary-gonadal axis when analyzing plasma concentrations. Therefore, we examined the relationship between the excretion of urinary LH and T in male Sprague-Dawley rats during exposure to increased gravity upon return to Earth following a 14-day spaceflight (n = 6) and by 12 days of centrifugation at 2g (n = 8). Excreted LH and T were elevated on the first 3 days postflight. Excreted T was elevated between Days 1 and 8 of centrifugation; however, excreted LH was reduced on Days 2 and 3 compared with control animals. Excreted LH and T were significantly correlated (R = 0.731 and 0.706, respectively) in postspaceflight and centrifuged animals. Correlation curves had similar slopes (0.0213 and 0.023, respectively), but different y-intercepts (-1.43 and 3.32, respectively). The sustained increase in excreted T during centrifugation suggests that the pituitary-gonadal axis in postspaceflight animals may adapt quicker to increased gravity. The upward shift in the correlation curve exhibited by the centrifuged animals suggests that the sensitivity of LH-induced T release is increased in these animals. The previous dissociation between plasma LH and T during altered gravity was not observed in the present study in which excreted LH and T were measured.

short duration↗

Plant adaptation to low atmospheric pressures: potential molecular responses

There is an increasing realization that it may be impossible to attain Earth normal atmospheric pressures in orbital, lunar, or Martian greenhouses, simply because the construction materials do not exist to meet the extraordinary constraints imposed by balancing high engineering requirements against high lift costs. This equation essentially dictates that NASA have in place the capability to grow plants at reduced atmospheric pressure. Yet current understanding of plant growth at low pressures is limited to just a few experiments and relatively rudimentary assessments of plant vigor and growth. The tools now exist, however, to make rapid progress toward understanding the fundamental nature of plant responses and adaptations to low pressures, and to develop strategies for mitigating detrimental effects by engineering the growth conditions or by engineering the plants themselves. The genomes of rice and the model plant Arabidopsis thaliana have recently been sequenced in their entirety, and public sector and commercial DNA chips are becoming available such that thousands of genes can be assayed at once. A fundamental understanding of plant responses and adaptation to low pressures can now be approached and translated into procedures and engineering considerations to enhance plant growth at low atmospheric pressures. In anticipation of such studies, we present here the background arguments supporting these contentions, as well as informed speculation about the kinds of molecular physiological responses that might be expected of plants in low-pressure environments.

NASA Discipline Plant Biology↗

Final Science Reports of the US Experiments Flown on the Russian Biosatellite Cosmos 2229

Cosmos 2229 was launched on December 29, 1992, containing a biological payload including two young male rhesus monkeys, insects, amphibians, and cell cultures. The biosatellite was launched from the Plesetsk Cosmodrome in Russia for a mission duration of 11.5 days. The major research objectives were: (1) Study of adaptive response mechanisms of mammals during flight; and (2) Study of physiological mechanisms underlying vestibular, motor system and brain function in primates during early and later adaptation phases. American scientists and their Russian collaborators conducted 11 experiments on this mission which included extensive preflight and postflight studies with rhesus monkeys. Biosamples and data were subsequently transferred to the United States. The U.S. responsibilities for this flight included the development of experiment protocols, the fabrication of some flight instrumentation and experiment-specific ground-based hardware, the conducting of preflight and postflight testing and the analysis of biospecimens and data for the U.S. experiments. A description of the Cosmos 2229 mission is presented in this report including preflight, on-orbit and postflight activities. The flight and ground-based bioinstrumentation which was developed by the U.S. and Russia is also described, along with the associated preflight testing ot the U.S. hardware. Final Science Reports for the experiments are also included.

Connolly, James P.↗

Space research activities during missions of the past

For over 30 yr, men and women have lived for various periods of time in a weightless (or free-fall) environment while orbiting the Earth. During these years, we have learned that humans function quite well for short periods of weightlessness, that is, for up to a little more than a year. Some space flight missions have provided physiologic data, including cardiovascular data, from the spacefarers. In fact, some missions have provided laboratories for systematic study of cardiovascular responses and adaptation to space flight. However, the opportunity to obtain physiologic data from people in space is a rarity. It is important to remember that the population sample sizes are small, other stresses may confound the effects of weightlessness, and in some situations the crewmembers are subjects for several experiments at the same time. Furthermore, comparison of cardiovascular data from space flight to data obtained on the ground is sometimes difficult because the subject's posture on the ground is not always reported; in a gravity environment, posture influences the hydrostatic gradient. This over view describes what we have learned about cardiovascular function during flight and after return to Earth.

NASA Discipline Cardiopulmonary↗

Erythropoietin withdrawal alters interactions between young red blood cells, splenic endothelial cells, and macrophages: an in vitro model of neocytolysis

BACKGROUND: We have described the rapid destruction of young red blood cells (neocytolysis) in astronauts adapting to microgravity, in polycythemic high altitude dwellers who descend to sea level, and in patients with kidney disorders. This destruction results from a decrease in erythropoietin (EPO) production. We hypothesized that such EPO withdrawal could trigger physiological changes in cells other than red cell precursors and possibly lead to the uptake and destruction of young red cells by altering endothelial cell-macrophage interactions, most likely occurring in the spleen. METHODS: We identified EPO receptors on human splenic endothelial cells (HSEC) and investigated the responses of these cells to EPO withdrawal. RESULTS: A monolayer of HSEC, unlike human endothelial cells from aorta, glomerulus, or umbilical vein, demonstrated an increase in permeability upon EPO withdrawal that was accompanied by unique morphological changes. When HSEC were cultured with monocyte-derived macrophages (but not when either cell type was cultured alone), EPO withdrawal induced an increased ingestion of young red cells by macrophages when compared with the constant presence or absence of EPO. CONCLUSIONS: HSEC may represent a unique cell type that is able to respond to EPO withdrawal by increasing permeability and interacting with phagocytic macrophages, which leads to neocytolysis.

NASA Discipline Cardiopulmonary↗

Enabling human exploration of space - A life sciences overview

In the transition from the short-duration missions of the Space Shuttle era to long-duration exploration missions, the health and safety of crewmembers must be ensured. The body undergoes many complex physiological changes as a result of its adaptation to a microgravity environment and U.S. and Soviet experiences have shown that time is required for readaptation to gravity. The consequences of these changes for the extended exploration missions envisioned for the future are unknown. A Mars mission may require crewmembers to spend many months in microgravity, and then work effectively in a one-third gravity environment. Other problems may arise when returning crewmembers must readapt to earth's gravity. Life Sciences activities are being planned to systematically address the physiological issues involved with long-term manned exploration missions, through ground-based studies and flight investigations on the Shuttle and Space Station Freedom. The areas of focus are artificial gravity, radiation, health care, and space human factors.

Gaiser, Karen K.↗

Unimpaired Neuro-Adaptive Plasticity in an Elderly Astronaut

Quantitative analyses of a 77 year old astronaut's balance control performances on a standardized test battery revealed few differences between his neuro-adaptive responses to space flight and those of a group of younger astronauts tested following missions of similar duration. This finding suggests that the physiological changes associated with age do not necessarily impair adaptive plasticity in the human following removal and subsequent reintroduction of gravity.

Paloski, William H.↗

Crew Factors in Flight Operations 7: Psychophysiological Responses to Overnight Cargo Operations

To document the psychophysiological effects of flying overnight cargo operations, 41 B-727 crew members (average age 38 yr) were monitored before, during, and after one of two typical 8-day trip patterns. During daytime layovers, the average sleep episode was 3 hr (41%) shorter than nighttime sleeps and was rated as lighter, less restorative, and poorer overall. Sleep was frequently split into several episodes and totaled 1.2 hr less per 24 hr than on pretrip days. Each trip pattern included a night off, which was an effective countermeasure against the accumulating sleep debt. The organization of sleep during daytime layovers reflected the interaction of duty timing with circadian physiology. The circadian temperature rhythm did not adapt completely to the inverted wake-rest schedule on duty days, being delayed by about 3 hr. Highest subjective fatigue and lowest activation occurred around the time of the temperature minimum. On duty days, reports of headaches increased by 400%, of congested nose by 200%, and of burning eyes by 900%. Crew members also reported eating more snacks. Compared with daytime short-haul air-transport operations, the overnight cargo trips included fewer duty and flight hours, and had longer layovers. Overnight cargo crews also averaged 5.4 yr younger than their daytime short-haul counterparts. On trips, both groups lost a comparable amount of sleep per 24 hr, but the overnight cargo crews had shorter individual sleep episodes and more broken sleep. These data clearly demonstrate that overnight cargo operations, like other night work, involve physiological disruption not found in comparable daytime operations.

Gander, Philippa H.↗

Microbial Responses to Microgravity and Other Low-Shear Environments

Microbial adaptation to environmental stimuli is essential for survival. While several of these stimuli have been studied in detail, recent studies have demonstrated an important role for a novel environmental parameter in which microgravity and the low fluid shear dynamics associated with microgravity globally regulate microbial gene expression, physiology, and pathogenesis. In addition to analyzing fundamental questions about microbial responses to spaceflight, these studies have demonstrated important applications for microbial responses to a ground-based, low-shear stress environment similar to that encountered during spaceflight. Moreover, the low-shear growth environment sensed by microbes during microgravity of spaceflight and during ground-based microgravity analogue culture is relevant to those encountered during their natural life cycles on Earth. While no mechanism has been clearly defined to explain how the mechanical force of fluid shear transmits intracellular signals to microbial cells at the molecular level, the fact that cross talk exists between microbial signal transduction systems holds intriguing possibilities that future studies might reveal common mechanotransduction themes between these systems and those used to sense and respond to low-shear stress and changes in gravitation forces. The study of microbial mechanotransduction may identify common conserved mechanisms used by cells to perceive changes in mechanical and/or physical forces, and it has the potential to provide valuable insight for understanding mechanosensing mechanisms in higher organisms. This review summarizes recent and future research trends aimed at understanding the dynamic effects of changes in the mechanical forces that occur in microgravity and other low-shear environments on a wide variety of important microbial parameters.

Review↗