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All sourcesNASA NTRSDOE OSTI
NASA NTRS1999

Non-Invasive Investigation of Bone Adaptation in Humans to Mechanical Loading

Experimental studies have identified peak cyclic forces, number of loading cycles, and loading rate as contributors to the regulation of bone metabolism. We have proposed a theoretical model that relates bone density to a mechanical stimulus derived from average daily cumulative peak cyclic 'effective' tissue stresses. In order to develop a non-invasive experimental model to test the theoretical model we need to: (1) monitor daily cumulative loading on a bone, (2) compute the internal stress state(s) resulting from the imposed loading, and (3) image volumetric bone density accurately, precisely, and reproducibly within small contiguous volumes throughout the bone. We have chosen the calcaneus (heel) as an experimental model bone site because it is loaded by ligament, tendon and joint contact forces in equilibrium with daily ground reaction forces that we can measure; it is a peripheral bone site and therefore more easily and accurately imaged with computed tomography; it is composed primarily of cancellous bone; and it is a relevant site for monitoring bone loss and adaptation in astronauts and the general population. This paper presents an overview of our recent advances in the areas of monitoring daily ground reaction forces, biomechanical modeling of the forces on the calcaneus during gait, mathematical modeling of calcaneal bone adaptation in response to cumulative daily activity, accurate and precise imaging of the calcaneus with quantitative computed tomography (QCT), and application to long duration space flight.

Whalen, R.↗
NASA NTRS2008

Effect of Vestibular Impairment on Cerebral Blood Flow Response to Dynamic Roll Tilt

Change to upright posture results in reductions in cerebral perfusion pressure due to hydrostatic pressure changes related to gravity. Since vestibular organs, specifically the otoliths, provide information on position relative to gravity, vestibular inputs may assist in adaptation to the upright posture. The goal of this study was to examine the effect of direct vestibular stimulation on cerebral blood flow (CBF). To examine the role of otolith inputs we screened 165 subjects for vestibular function and classified subjects as either normal or impaired based on ocular torsion. Ocular torsion, an indication of otolith function, was assessed during sinusoidal roll tilt of 20 degrees at 0.01 Hz (100 sec per cycle). Subjects with torsion one SD below the mean were classified as impaired while subjects one SD above the mean were considered normal. During one session subjects were placed in a chair that was sinusoidally rotated 25 degrees in the roll plane at five frequencies: 0.25 & 0.125 Hz for 80 sec, 0.0625 Hz for 160 sec and 0.03125 Hz and 0.015625 Hz for 320 sec. During testing, CBF (transcranial Doppler), blood pressure (Finapres), and end tidal CO2 (Puritan Bennet) were measured continuously. Ocular torsion was assessed from infrared images of the eyes. All rotations were done in the dark with subjects fixated on a red LED directly at the center of rotation. In the normal group, dynamic tilt resulted in significant changes in both blood pressure and cerebral blood flow velocity that was related to the frequency of stimulus. In contrast the impaired group did not show similar patterns. As expected normal subjects demonstrated significant ocular torsion that was related to stimulus frequency while impaired subjects had minimal changes. These data suggest that vestibular inputs have direct effects on cerebral blood flow regulation during dynamic tilt. Supported by NASA.

Serrador, J. M.↗
NASA NTRS1991

Reassessment of area postrema's role in motion sickness and conditioned taste aversion

On the basis of classical studies on the role of the area psotrema (AP) in motion-induced emesis it was generally accepted that the AP is an essential structure for the production of vomiting in response to motion. However, in more recent studies it has been demonstrated that vomiting induced by motion can still occur in animals in which the AP has been destroyed bilaterally. It was inferred from some of these more recent studies that the AP plays no role in motion-induced emesis. From the standpoint of the current understanding of central nervous system (CNS) plasticity, redundancy, remodeling, unmasking, regeneration, and recovery of function, however, it is important to realize the limitations of using ablation procedures to determine the functional role of a given neural structure in a highly integrated, adaptable central nervous system (CNS). For example, the results of our recent investigations in cat and squirrel monkey on the role of the AP in emesis and conditioned taste aversion induced by motion indicate that while AP lesions do not prevent motion-induced emesis when animals are tested 30 days or more after surgery, the lesions do change the latency to emesis. Thus, contradictory findings from lesion studies must be evaluated not only in terms of species difference, differences in lesioning techniques and extent of lesions, and in stimulus parameters, but also in terms of duration of the recovery period, during which significant recovery of function may take place. In our judgment, inadequate consideration of the foregoing factors could lead to erroneous inferences about given structure's role in the behavior of the intact, nonablated animal.

N G Daunton↗
NASA NTRSDate not supplied

Single-Cell mRNA Sequencing of Bone Marrow Mesenchymal Stem Cell Lineages Shows that Gravity Mechanical Loading Regulates Regenerative Osteogenesis in a CDKN1A-Dependent Manner

Weight-bearing at 1g is increasingly recognized as an important stimulus driving stem cell-based tissue regenerative homeostasis and health. In the bone marrow compartment, mechanical stimuli from gravity can drive osteogenesis from mesenchymal progenitors and hematopoiesis from hematopoietic progenitors. Conversely, hematopoietic monocyte/macrophage progenitors can also be driven into osteoclastogenesis by mechanical unloading in microgravity. Although the cell cycle inhibitor gene Cdkn1a is known to be upregulated in microgravity, the great diversity of stem cell lineages and stages of differentiation in bone marrow, make it difficult to attribute gene expression changes to specific cell types. To overcome this limitation, we used single-cell mRNAseq analyses with both in-vitro and in-vivo strategies, including in-vitro stretch of marrow osteogenic cultures, and in-vivo hindlimb unloading, running wheels, treadmills, and exposure to real microgravity. For in-vitro studies marrow osteoprogenitors from WT and Cdkn1a-null mice were subjected to substrate stretch during osteogenesis, with static cultures simulating unloading. From those experiments we determined Cdkn1a-null cells transition more readily from progenitors to early osteoblasts and further to mineralizing osteoblasts. Additionally, single-cell mRNAseq resolution identified Cdkn1a suppression in response to mechanical loading predominantly in early proliferative osteoblasts rather than progenitor or mineralizing osteoblasts. These findings support the hypothesis that CDKN1A plays a mechano-reversible, anti-proliferative role during bone regenerative osteogenesis. To investigate in-vivo mechanoregulation of osteogenesis, mice were flown in microgravity during the Rodent Research-10 experiment comparing wildtype and Cdkn1a-null mice kept on ISS for 30 days. The mice were euthanized and dissected on-orbit in microgravity and femur bone marrow was collected using a custom sample isolation and live-cell cryo-preservation technique allowing for single cell transcriptomics post-flight processing. Ground control experiments with hindlimb-unloading, running wheels, and treadmills, were also conducted for a comprehensive analysis of mechanical loading effects on the regenerative health of bone tissue and its marrow stem cell progenitor lineages.

single cell sequencing↗
NASA NTRS1986

The Central Nervous Connections Involved in the Vomiting Reflex

The vomiting reflex may be elicited by a number of different types or classes of stimuli involving many varieties of receptor structures and considerable diversity in afferent pathways and central connections. Central relay or mediating structures thus may vary widely according to the type of initial emetic stimulus. The emetic circuits which have been most completely delineated to date are probably those in which the Chemoreceptor Trigger Zone (CTZ) in the Area Postrema (AP) functions as a key mediating structure. Even in this system, however, there are large gaps in our knowledge of the nerve tracts and central nervous connections involved. Knowledge of most other emetic circuits subserving the emetic reflex resulting from many diverse types of stimuli such, for example, as emotional stress (e.g. psychogenic vomiting, Wruble et al. 1982), pain (e.g. testicular trauma), and chemical or mechanical irritation of the gastrointestinal tract or urinary tract is quite incomplete at this time, thus precluding any very adequate description of their central connections at present. One physiological system, however, which has received considerable attention recently in relation to the vomiting reflex elicited by motion stimuli is the vestibular system. Due to the paucity of data on central nervous connections of several or the non-vestibular types of emetic stimuli cited above, we will devote most of our attention in this brief review to the central connections of the vestibular system which seem likely to be involved in the vomiting response to motion stimuli. However, the latter part of the review will be concerned with the concept of the reticular vomiting centre in relation to the ParviCellular Reticular Formation (PCRF), and will thus probably pertain to all of the many classes of emetic stimuli since it will address the question of the final common emetic pathway.

Brizzee, K. R.↗
NASA NTRS1999

Ventilatory responses to dynamic exercise elicited by intramuscular sensors

PURPOSE: Eight subjects, aged 27.0+/-1.6 yr, performed incremental workload cycling to investigate the contribution of skeletal muscle mechano- and metaboreceptors to ventilatory control during dynamic exercise. METHODS: Each subject performed four bouts of exercise: exercise with no intervention (CON); exercise with bilateral thigh cuffs inflated to 90 mm Hg (CUFF); exercise with application of lower-body positive pressure (LBPP) to 45 torr (PP); and exercise with 90 mm Hg thigh cuff inflation and 45 torr LBPP (CUFF+PP). Ventilatory responses and pulmonary gas exchange variables were collected breath-by-breath with concomitant measurement of leg intramuscular pressure. RESULTS: Ventilation (VE) was significantly elevated from CON during PP and CUFF+PP at workloads corresponding to > or = 60% CON peak oxygen uptake (VO2peak) and during CUFF at workloads > or = 80% CON VO2peak, P < 0.05. The VO2 at which ventilatory threshold occurred was significantly reduced from CON (2.17+/-0.28 L x min(-1)) to 1.60+/-0.19 L x min(-1), 1.45+/-0.15 L x min(-1), and 1.15+/-0.11 L x min(-1) during CUFF, PP, and CUFF+PP, respectively. The slope of the linear regression describing the VE/CO2 output relationship was increased from CON by approximately 22% during CUFF, 40% during PP, and 41% during CUFF+PP. CONCLUSIONS: As intramuscular pressure was significantly elevated immediately upon application of LBPP during PP and CUFF+PP without a concomitant increase in VE, it seems unlikely that LBPP-induced increases in VE can be attributed to activation of the mechanoreflex. These findings suggest that LBPP-induced reductions in perfusion pressure and decreases in venous outflow resulting from inflation of bilateral thigh cuffs may generate a metabolite sensitive intramuscular ventilatory stimulus.

Non-NASA Center↗
NASA NTRS1997

Modulation of statolith mass and grouping in white clover (Trifolium repens) growth in 1-g, microgravity and on the clinostat

Current models of gravity perception in higher plants focus on the buoyant weight of starch-filled amyloplasts as the initial gravity signal susceptor (statolith). However, no tests have yet determined if statolith mass is regulated to increase or decrease gravity stimulus to the plant. To this end, the root caps of white clover (Trifolium repens) grown in three gravity environments with three different levels of gravity stimulation have been examined: (i) 1-g control with normal static gravistimulation, (ii) on a slow clinostat with constant gravistimulation, and (iii) in the stimulus-free microgravity aboard the Space Shuttle. Seedlings were germinated and grown in the BioServe Fluid Processing Apparatus and root cap structure was examined at both light and electron microscopic levels, including three-dimensional cell reconstruction from serial sections. Quantitative analysis of the electron micrographs demonstrated that the starch content of amyloplasts varied with seedling age but not gravity condition. It was also discovered that, unlike in starch storage amyloplasts, all of the starch granules of statolith amyloplasts were encompassed by a fine filamentous, ribosome-excluding matrix. From light micrographic 3-D cell reconstructions, the absolute volume, number, and positional relationships between amyloplasts showed (i) that individual amyloplast volume increased in microgravity but remained constant in seedlings grown for up to three days on the clinostat, (ii) the number of amyloplasts per cell remained unchanged in microgravity but decreased on the clinostat, and (iii) the three-dimensional positions of amyloplasts were not random. Instead amyloplasts in microgravity were grouped near the cell centers while those from the clinostat appeared more dispersed. Taken together, these observations suggest that changing gravity stimulation can elicit feedback control over statolith mass by changing the size, number, and grouping of amyloplasts. These results support the starch-statolith theory of graviperception in higher plants and add to current models with a new feedback control loop as a mechanism for modulation of statolith responsiveness to inertial acceleration.

Flight Experiment↗
NASA NTRS1994

Interaction of chronic reatine depletion and muscle unloading effects on postural and locomotor muscles

In this study, creatine depletion was induced separately and in combination with non-weight-bearing activity to determine if the response to lowering this metabolite would counter the MHC transitions expected from non-weight-bearing. Creatine depletion was induced by feeding rats a diet supplemented with the creatine analogue beta-guanidinopropionic acid (beta-GPA). Animals were fed a diet containing the creatine analogue for 68 days. Hindlimb non-weight-bearing in BS and NS animals was accomplished by tail suspension for the final 30 days of this period. Beta-GPA feeding lowered the creatine content of muscles sampled by 65%. Creatine depletion resulted in a 16% increase in citrate synthase activity in the soleus (SOL) and a 24% increase in the plantaris (PLN). In two postural muscles, the SOL and vastus intermedius (VI), tail suspension resulted in large decreases in the type I MHC expression and increases in type IIx and IIb MHCs. In two locomotor muscles, the PLN and medial gastrocnemius, type I MHC declined and type IIb increased with suspension. Creatine depletion did not prevent the suspension-induced decline in type I MHC in any of these muscles. The increase in type IIb MHC was either prevented or reduced by creatine depletion before and during suspension in the SOL, VI, and PLN. Creatine depletion alone resulted in small increases in type I and IIa MHCs in the two locomotor muscles, but it had no effect on the MHC profile of the postural muscles studied. These results indicate that the mechanical signal generated by the hindlimb non-weight-bearing state dominated over the metabolic stimulus of creatine depletion with respect to the primary adaptation involving a reduction in type I MHC.

Adams, Gregory R.↗
NASA NTRS1999

Neurovestibular Session Summary

Oman - The early mission operational problems caused by space motion sickness have been largely resolved in recent years. This has been achieved by appropriate timeline adjustments, voluntary head movement restriction, and judicious use of promethazine. Crew members now simply accept that some symptoms "come with the job," and usually last only a few days. But as more people have flown longer flights, we've seen cases of space sickness and inversion illusion that take several weeks to resolve. Visual reorientation illusions continue throughout long flights, and occasionally cause difficulties. EVA astronauts sometimes suddenly fear they will fall out of the payload bay or off of the RMS or Strella arms. Orientation and navigation in three dimensions in the MIR station reportedly does not come naturally, because modules have different visual verticals. It is clear that the neurovestibular problems of spaceflight have not disappeared. After return to Earth, many crew members are disoriented and ataxic in the first hour after return, and require assistance leaving the vehicle, Flight surgeons say that the longer the mission, the stronger the aftereffects, certain of which last for weeks. We do not yet know how to predict who will be afflicted. Looking ahead to 3-4 month long voyages to Mars, it seems obvious that if cruise is in O-G, the crew may encounter neurovestibular problems on arrival. Artificial G may be broadly effective as a countermeasure for many of the physiological changes of spaceflight, but from the neurovestibular perspective, it is a double-edged sword. We know that the Coriolis stimulus resulting from rotation is potentially disorienting and nauseogenic. But we don't yet know how much artificial G will be enough, nor how successfully people can adapt to a specific angular velocity and hypo G level. Development of countermeasures remains a big challenge for our neurovestibular community. Maintaining an interdisciplinary perspective is important. Three examples were presented at this meeting: 1) Transgenic animal experiments suggest that in addition to the light illumination cycle, vestibular inputs may also serve as an important input to the circadian system. 2) Radiation can cause important CNS effects in animals, including loss of spatial memory. 3) As described in our session, otolith inputs may contribute to cardiovascular regulation of orthostatic tolerance. Over the past three days, we've all enjoyed catching up with old friends, and making many new ones. On behalf of my colleagues, I want to thank Al Coats and the USRA DSLS staff for the great job they did in running this meeting. And keeping the emphasis on fun. And also my Co- Chair, Mal Cohen, who had more stamina than many of us, despite major surgery only three weeks ago. Mal and I have written a few lines describing each of the seventeen papers in our session, to give you a quick over-view, and as a guide to the full abstracts, We have grouped them under five themes: preflight and inflight countermeasurements, postlanding posture and locomotion deficits: assessment and prediction, adaptive processes, relationships among physical simuli, perceptions, and eye movements, vestibular contribution to human autonomic responses, and implications and recommendations.

Oman, Charles↗
NASA NTRS2002

Smooth Pursuit Saccade Amplitude Modulation During Exposure to Microgravity

Russian investigators have reported changes in pursuit tracking of a vertically moving point stimulus during space flight. Early in microgravity, changes were manifested by decreased eye movement amplitude (undershooting) and the appearance of correction saccades. As the flight progressed, pursuit of the moving point stimulus deteriorated while associated saccadic movements were unchanged. Immediately postflight there was an improved execution of active head movements indicating that the deficiencies in pursuit function noted in microgravity may be of central origin. In contrast, tests of two cosmonauts showed that horizontal and vertical smooth pursuit were unchanged inflight. However, results of corresponding saccadic tasks showed a tendency toward the overshooting of a horizontal target early inflight with high accuracy developing later inflight, accompanied by an increased saccade velocity and a trend toward decreased saccade latency. Based on these equivocal results, we have further investigated the effects of space flight on the smooth pursuit mechanism during and after short duration flight, and postflight on returning MIR crewmembers. Sinusoidal target movement was presented horizontally at frequencies of 0.33 and 1.0 Hz. Subjects were asked to perform two trials for each stimulus combination: (1) moving eyes-only (EO) and (2) moving eyes and head (EH) with the target motion. Peak amplitude was 30 deg for 0.33 Hz trials and 15 deg for the 1.0 Hz trials. The relationship between saccade amplitude and peak velocity were plotted as a main sequence for each phase of flight, and linear regression analysis allowed us to determine the slope of each main sequence plot. The linear slopes were then combined for each flight phase for each individual subject. The main sequence for both EO and EH trials at both the 0.33 and 1.0 Hz frequencies during flight for the short duration flyers showed a reduction in saccade velocity and amplitude when compared to the preflight main sequence . This difference in the regression slopes between flight phase, head/eye condition (EO or EH), and pursuit target frequency was observed across all subjects (statistically significant at the p<0.02, df= 2). It is interesting to note that postflight for the short duration flyers there was an immediate recovery to the preflight main sequence across all trials. There were no significant differences observed between the preflight slopes for either head movement condition (EO vs. EH). When the immediate postflight (R+O) regression slopes were compared with the preflight slopes, there was a tendency (not significant) for both saccade amplitude and peak velocity to increase during the postflight testing. This tendency had vanished by R+ 1. Of particular interest was the redistribution of saccades during the latter stages of the flight and immediately postflight in the EO condition. At the 1.0 Hz frequency the saccades tended to be clustered near the lowest target velocity. It was also interesting to note that gaze performance (eye in skull + head in space) was consistently better during the EH condition; a finding also observed by our Russian colleagues. As the results of the long duration flight become available we expect that they will not only show that postflight effects will be similar to those observed during the short duration flights, but will also last for a greater period of time following flight. It is not clear what mechanism is responsible for the decreased peak saccadic velocity during flight unless the change is related to the control of retinal slip. For example, it is possible that saccades will tend to initially undershoot their targets by a small percentage and these saccades are then followed, if vision is available, by a small augmenting corrective saccade. It has been postulated that the functional significance of this undershooting tendency is to maintain the spatial representation of the target on the same side of the fovea (as opposedo racing across the fovea) and hence in the same cerebral hemisphere that initiated the primary saccade thus minimizing delays caused by an intra-hemispheric transfer of information . One could also speculate that with saccade velocities greater than normal, additional corrective saccades would be required to bring the target back on the fovea. A less plausible explanation of our findings could be fatigue. Yet it seems unlikely that our subjects would show lower velocities on all inflight test days while showing increased saccade velocities immediately following space flight where fatigue is usually the greatest. Finally, the redistribution effect noted late inflight is likely caused by adaptive changes. Overall, corrective saccades appeared to be used in maintaining gaze on target; reducing retinal slip and assisting space travelers in maintaining clear vision throughout the different phases of the space flight.

Reschke, M. F.↗