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PubMed2003

Towards new tests of strong-field gravity with measurements of surface atomic line redshifts from neutron stars.

In contrast to gravity in the weak-field regime, which has been subject to numerous experimental tests, gravity in the strong-field regime is largely unconstrained by observations. We show that gravity theories that pass solar system tests, but that diverge from general relativity in the strong-field regime, predict neutron stars with significantly different properties than their general relativistic counterparts. The range of redshfits of surface atomic lines predicted by such theories is significantly wider than the uncertainty introduced by our lack of knowledge of the equation of state of ultradense matter. Measurements of such lines with x-ray observatories can thus put new constraints on strong-field gravity.

Journal Article↗
PubMed2005

Brane-induced-gravity shock waves.

We construct exact gravitational field solutions for a relativistic particle localized on a tensional brane in brane-induced gravity. They are a generalization of gravitational shock waves in 4D de Sitter space. We provide the metrics for both the normal branch and the self-inflating branch Dvali-Gabadadze-Porrati brane worlds, and compare them to the 4D Einstein gravity solution and to the case when gravity resides only in the 5D bulk, without any brane-localized curvature terms. At short distances the wave profile looks the same as in four dimensions. The corrections appear only far from the source, where they differ from the long distance corrections in 4D de Sitter space. We also discover a new nonperturbative channel for energy emission into the bulk from the self-inflating [corrected] branch, when gravity is modified at the de Sitter radius.

Journal Article↗
PubMed1992

Ionic Current Changes Associated with the Gravity-Induced Bending Response in Roots of Zea mays L.

A vibrating probe was used to measure the changes in ionic currents around gravistimulated roots of Zea mays L. in an effort to determine whether these currents are involved in stimulus transduction from the root cap to the elongation zone. We did not observe a migration of the previously reported auxin-insensitive current efflux associated with gravity sensing (T. Björkman, A.C. Leopold [1987] Plant Physiol 84:841-846) back from the root cap. Instead, beginning 10 to 15 min after gravistimulation, an asymmetry in current developed simultaneously along the root around the meristem and apical regions of the elongation zone. This asymmetry comprised a proton efflux from the upper surface, which was superimposed on the symmetrical pattern around the vertical root. The gravity-induced proton efflux was inhibited by the application of the auxin transport inhibitor, 2,3,5-triiodobenzoic acid, whereas the calcium channel blocker, lanthanum, had little effect. Because the onset of the gravity-induced current asymmetry coincided both spatially and temporally with the onset of the differential growth response, we suggest that this current efflux may result from auxin-requiring acid-growth phenomena in the upper root tissue. The implications of this simultaneous onset of both proton efflux and elongation for theories about gravity stimulus transduction are discussed.

Journal Article↗
PubMed2004

Interactions between auxin transport and the actin cytoskeleton in developmental polarity of Fucus distichus embryos in response to light and gravity.

Land plants orient their growth relative to light and gravity through complex mechanisms that require auxin redistribution. Embryos of brown algae use similar environmental stimuli to orient their developmental polarity. These studies of the brown algae Fucus distichus examined whether auxin and auxin transport are also required during polarization in early embryos and to orient growth in already developed tissues. These embryos polarize with the gravity vector in the absence of a light cue. The auxin, indole-3-acetic acid (IAA), and auxin efflux inhibitors, such as naphthylphthalamic acid (NPA), reduced environmental polarization in response to gravity and light vectors. Young rhizoids are negatively phototropic, and NPA also inhibits rhizoid phototropism. The effect of IAA and NPA on gravity and photopolarization is maximal within 2.5 to 4.5 h after fertilization (AF). Over the first 6 h AF, auxin transport is relatively constant, suggesting that developmentally controlled sensitivity to auxin determines the narrow window during which NPA and IAA reduce environmental polarization. Actin patches were formed during the first hour AF and began to photolocalize within 3 h, coinciding with the time of NPA and IAA action. Treatment with NPA reduced the polar localization of actin patches but not patch formation. Latrunculin B prevented environmental polarization in a time frame that overlaps the formation of actin patches and IAA and NPA action. Latrunculin B also altered auxin transport. Together, these results indicate a role for auxin in the orientation of developmental polarity and suggest interactions between the actin cytoskeleton and auxin transport in F. distichus embryos.

Actins↗
PubMed2004

The fast and transient transcriptional network of gravity and mechanical stimulation in the Arabidopsis root apex.

Plant root growth is affected by both gravity and mechanical stimulation (Massa GD, Gilroy S [2003] Plant J 33: 435-445). A coordinated response to both stimuli requires specific and common elements. To delineate the transcriptional response mechanisms, we carried out whole-genome microarray analysis of Arabidopsis root apices after gravity stimulation (reorientation) and mechanical stimulation and monitored transcript levels of 22,744 genes in a time course during the first hour after either stimulus. Rapid, transient changes in the relative abundance of specific transcripts occurred in response to gravity or mechanical stimulation, and these transcript level changes reveal clusters of coordinated events. Transcriptional regulation occurs in the root apices within less than 2 min after either stimulus. We identified genes responding specifically to each stimulus as well as transcripts regulated in both signal transduction pathways. Several unknown genes were specifically induced only during gravitropic stimulation (gravity induced genes). We also analyzed the network of transcriptional regulation during the early stages of gravitropism and mechanical stimulation.

Arabidopsis↗
PubMed1998

Mapping the functional roles of cap cells in the response of Arabidopsis primary roots to gravity.

The cap is widely accepted to be the site of gravity sensing in roots because removal of the cap abolishes root curvature. Circumstantial evidence favors the columella cells as the gravisensory cells because amyloplasts (and often other cellular components) are polarized with respect to the gravity vector. However, there has been no functional confirmation of their role. To address this problem, we used laser ablation to remove defined cells in the cap of Arabidopsis primary roots and quantified the response of the roots to gravity using three parameters: time course of curvature, presentation time, and deviation from vertical growth. Ablation of the peripheral cap cells and tip cells did not alter root curvature. Ablation of the innermost columella cells caused the strongest inhibitory effect on root curvature without affecting growth rates. Many of these roots deviated significantly from vertical growth and had a presentation time 6-fold longer than the controls. Among the two inner columella stories, the central cells of story 2 contributed the most to root gravitropism. These cells also exhibited the largest amyloplast sedimentation velocities. Therefore, these results are consistent with the starch-statolith sedimentation hypothesis for gravity sensing.

Arabidopsis↗
PubMed2001

Cytoplasmic pH dynamics in maize pulvinal cells induced by gravity vector changes.

In maize (Zea mays) and other grasses, changes in orientation of stems are perceived by pulvinal tissue, which responds to the stimulus by differential growth resulting in upward bending of the stem. The amyloplast-containing bundle sheath cells are the sites of gravity perception, although the initial steps of gravity perception and transmission remain unclear. In columella cells of Arabidopsis roots, we previously found that cytoplasmic pH (pH(c)) is a mediator in early gravitropic signaling (A.C. Scott, N.S. Allen [1999] Plant Physiol 121: 1291-1298). The question arises whether pH(c) has a more general role in signaling gravity vector changes. Using confocal ratiometric imaging and the fluorescent pH indicator carboxy seminaphtorhodafluor acetoxymethyl ester acetate, we measured pH(c) in the cells composing the maize pulvinus. When stem slices were gravistimulated and imaged on a horizontally mounted confocal microscope, pH(c) changes were only apparent within the bundle sheath cells, and not in the parenchyma cells. After turning, cytoplasmic acidification was observed at the sides of the cells, whereas the cytoplasm at the base of the cells where plastids slowly accumulated became more basic. These changes were most apparent in cells exhibiting net amyloplast sedimentation. Parenchyma cells and isolated bundle sheath cells did not show any gravity-induced pH(c) changes although all cell types responded to external stimuli in the predicted way: Propionic acid and auxin treatments induced acidification, whereas raising the external pH caused alkalinization. The results suggest that pH(c) has an important role in the early signaling pathways of maize stem gravitropism.

Cytoplasm↗
PubMed1966

Orientation of wheat seedling organs in relation to gravity.

Seedlings of wheat (Triticum aestivum L.) were grown in special holders that permitted the coleoptile and early roots to develop in moist air. The orientation of the organs of seedlings erect to gravity was compared with that of organs produced on a horizontal clinostat. Orientation was described by the angular position of each organ tip with reference to the axis of the embryo. Comparative tests were also made with barley, rye, and oat seedlings.The coleoptile of all species developed curvatures in 3 dimensions when geotropic responses were eliminated. The primary root was not precise in its positive geotropism. Seedlings grew on clinostats with much greater variations in the lateral orientation of the central root and with a tendency for it to curve away from the endosperm to a greater degree than in erect seedlings.The symmetry of root system in wheat was found to depend on a specific mechanism. Under the influence of gravity the earliest lateral roots were oriented in a plane at characteristic angles of about 57.5 degrees with the ideal primary root. The corresponding angles for lateral roots growing on clinostats were greater by about 47.5 degrees as a result of epinasty not previously reported in roots. This force also appeared to be active in the seminal roots of barley and rye but not of oats.The curvatures in coleoptiles grown without the directional effects of gravity correspond to the results of growth imbalance in Coleus stems in the absence of lateral transport of their auxin by gravity. Root epinasty appears to be based on auxin imbalance. Curvatures in the primary root are also interpreted as results of asymmetrical distribution of growth hormone.

Journal Article↗
PubMed2001

Genetic and chemical reductions in protein phosphatase activity alter auxin transport, gravity response, and lateral root growth.

Auxin transport is required for important growth and developmental processes in plants, including gravity response and lateral root growth. Several lines of evidence suggest that reversible protein phosphorylation regulates auxin transport. Arabidopsis rcn1 mutant seedlings exhibit reduced protein phosphatase 2A activity and defects in differential cell elongation. Here we report that reduced phosphatase activity alters auxin transport and dependent physiological processes in the seedling root. Root basipetal transport was increased in rcn1 or phosphatase inhibitor-treated seedlings but showed normal sensitivity to the auxin transport inhibitor naphthylphthalamic acid (NPA). Phosphatase inhibition reduced root gravity response and delayed the establishment of differential auxin-induced gene expression across a gravity-stimulated root tip. An NPA treatment that reduced basipetal transport in rcn1 and cantharidin-treated wild-type plants also restored a normal gravity response and asymmetric auxin-induced gene expression, indicating that increased basipetal auxin transport impedes gravitropism. Increased auxin transport in rcn1 or phosphatase inhibitor-treated seedlings did not require the AGR1/EIR1/PIN2/WAV6 or AUX1 gene products. In contrast to basipetal transport, root acropetal transport was normal in phosphatase-inhibited seedlings in the absence of NPA, although it showed reduced NPA sensitivity. Lateral root growth also exhibited reduced NPA sensitivity in rcn1 seedlings, consistent with acropetal transport controlling lateral root growth. These results support the role of protein phosphorylation in regulating auxin transport and suggest that the acropetal and basipetal auxin transport streams are differentially regulated.

Arabidopsis↗
PubMed2002

Physical aspects of protein crystal growth investigated with the Advanced Protein Crystallization Facility in reduced-gravity environments.

The physicochemical aspects of protein crystallization in reduced-gravity environments ( micro g) have been investigated with the Advanced Protein Crystallization Facility during six space missions. This review summarizes the results, dealing with the mechanisms of nucleation and crystal growth and with the quality of the crystals that were obtained under reduced gravity as well as under normal gravity on earth. Statistical analyses of the experimental data strongly support the fact that micro g has a positive effect on crystallization and on crystal quality. A comparison of experiments and theories of protein crystallization in reduced-gravity environments is presented. Recommendations for improving the performance of protein crystallization experiments in micro g and on earth are discussed.

Animals↗
PubMedDate not supplied

Delineating depth to bedrock beneath shallow unconfined aquifers: a gravity transect across the Palmer River Basin.

In view of the increasing demand on ground water supplies in the northeastern United States, it is imperative to develop appropriate methods to geophysically characterize the most widely used sources of ground water in the region: shallow unconfined aquifers consisting of well-sorted, stratified glacial deposits laid down in bedrock valleys and channels. The gravity method, despite its proven value in delineating buried bedrock valleys elsewhere, is seldom used by geophysical contractors in this region. To demonstrate the method's effectiveness for evaluating such aquifers, a pilot study was undertaken in the Palmer River Basin in southeastern Massachusetts. Because bedrock is so shallow beneath this aquifer (maximum depth is 30 m), the depth-integrated mass deficiency of the overlying unconsolidated material was small, so that the observed gravity anomaly was on the order of 1 milligal (mGal) or less. Thus data uncertainties were significant. Moreover, unlike previous gravity studies elsewhere, we had no a priori information on the density of the sediment. Under such circumstances, it is essential to include model constraints and weighted least-squares in the inversion procedure. Among the model constraints were water table configuration, bedrock outcrops, and depth to bedrock from five water wells. Our procedure allowed us to delineate depth to bedrock along a 3.5 km profile with a confidence interval of 1.8 m at a nominal depth of 17 m. Moreover, we obtained a porosity estimate in the range of 39% to 44%. Thus the gravity method, with appropriate refinements, is an effective tool for the reconnaissance of shallow unconfined aquifers.

Environmental Monitoring↗
PubMed2005

Effect of gravity on liquid plug transport through an airway bifurcation model.

Many medical therapies require liquid plugs to be instilled into and delivered throughout the pulmonary airways. Improving these treatments requires a better understanding of how liquid distributes throughout these airways. In this study, gravitational and surface mechanisms determining the distribution of instilled liquids are examined experimentally using a bench-top model of a symmetrically bifurcating airway. A liquid plug was instilled into the parent tube and driven through the bifurcation by a syringe pump. The effect of gravity was adjusted by changing the roll angle (phi) and pitch angle (gamma) of the bifurcation (phi = gamma =0 deg was isogravitational). Phi determines the relative gravitational orientation of the two daughter tubes: when phi not equal to 0 deg, one daughter tube was lower (gravitationally favored) compared to the other. Gamma determines the component of gravity acting along the axial direction of the parent tube: when gamma not equal to 0 deg, a nonzero component of gravity acts along the axial direction of the parent tube. A splitting ratio Rs, is defined as the ratio of the liquid volume in the upper daughter to the lower just after plug splitting. We measured the splitting ratio, Rs, as a function of: the parent-tube capillary number (Cap); the Bond number (Bo); phi; gamma; and the presence of pre-existing plugs initially blocking either daughter tube. A critical capillary number (Cac) was found to exist below which no liquid entered the upper daughter (Rs = 0), and above which Rs increased and leveled off with Cap. Cac increased while Rs decreased with increasing phi, gamma, and Bo for blocked and unblocked cases at a given Cap > Ca,. Compared to the nonblockage cases, Rs decreased (increased) at a given Cap while Cac increased (decreased) with an upper (lower) liquid blockage. More liquid entered the unblocked daughter with a blockage in one daughter tube, and this effect was larger with larger gravity effect. A simple theoretical model that predicts Rs and Cac is in qualitative agreement with the experiments over a wide range of parameters.

Biological Transport↗
PubMed1999

Magma intrusion beneath long valley caldera confirmed by temporal changes in gravity

Precise relative gravity measurements conducted in Long Valley (California) in 1982 and 1998 reveal a decrease in gravity of as much as -107 +/- 6 microgals (1 microgal = 10(-8) meters per square second) centered on the uplifting resurgent dome. A positive residual gravity change of up to 64 +/- 15 microgals was found after correcting for the effects of uplift and water table fluctuations. Assuming a point source of intrusion, the density of the intruding material is 2.7 x 10(3) to 4.1 x 10(3) kilograms per cubic meter at 95 percent confidence. The gravity results require intrusion of silicate magma and exclude in situ thermal expansion or pressurization of the hydrothermal system as the cause of uplift and seismicity.

Journal Article↗
PubMed1979

Radiographic comparison of human lung shape during normal gravity and weightlessness.

Human lung shape was measured during zero gravity (0 G) to decide whether the normal vertical regional differences in ventilation are due directly to distortion of the elastic lung by its own weight, or instead, due indirectly to the effect of gravity on the shape of the rib cage and diaphragm. This was important because we previously established that weightlessness virtually abolishes the normal topographical inequality of ventilation (J. Appl. Physiol.: Respirat. Environ. Exercise Physiol. 45: 987-998, 1978). Chest radiographs were made after 10 s of a weightless flight trajectory aboard a NASA-Ames Research Center Learjet in both posterior-anterior and left lateral projections on five seated volunteers at residual volume, functional residual capacity, and total lung capacity. Lung shape was assessed by measuring lung heights and widths in upper, middle, and lower lung regions. We found no significant differences between any of the normal gravity (1 G) and o G measurements, although there was a slight tendency for the lung to become shorter and wider at o G (mean changes generally less than 3% or about 0.5 cm). By contrast, Grassino et al. (J. Appl. Physiol. 39: 997-1003, 1975) found no change in the vertical distribution of ventilation after voluntarily changing lung dimensions by more than 1 cm by moving the abdomen in or out. We conclude that gravity produces the topographical distribution of ventilation in the upright human lung by distorting the elastic lung tissue within the chest rather than by altering the shape of the rib cage and diaphragm.

Humans↗
PubMed1987

No gravity-independent gradient of blood flow distribution in dog lung.

The existence of a major gravity-independent gradient of blood flow in lungs has recently been described based on single photon emission computed tomography after intravenous injection of radioactively labeled macroaggregates. We wanted to test this hypothesis of a major gravity-independent gradient in lung blood flow in experiments with direct measurement of macroaggregate distribution in the dog lung. In six anesthetized (4 prone spontaneously breathing, 2 mechanically ventilated) dogs we injected 111In-labeled albumin macroaggregates intravenously. We killed the dogs, removed, inflated, and froze the lower lobes. We sliced the lobes 1 cm thick and made gamma camera images of the slices. We then cut three or four slices in each lobe into two or three concentric layers and measured the radioactivity per gram of tissue in a well-type gamma counter. In three of the dogs we also labeled the red cells (99mTc) so that blood volume in each sample could be determined. The gamma camera images were acquired on a 64 X 64 matrix with 4 X 4 mm pixels. On the numeric printouts from the individual slices we made two or three concentric layers and calculated activity per pixel in each layer. Neither by the well counting nor by the pixel analysis of the gamma scans did we detect any gravity-independent distribution of blood flow. With the well counting the distribution was the same whether macroaggregate activity was expressed per gram of tissue or per gram of blood-free tissue. We conclude that by direct measurements no major gravity-independent gradient of pulmonary blood flow can be detected in dog lungs.

Animals↗
PubMed2000

Altered gravity downregulates aquaporin-1 protein expression in choroid plexus.

Aquaporin-1 (AQP1) is a water channel expressed abundantly at the apical pole of choroidal epithelial cells. The protein expression was quantified by immunocytochemistry and confocal microscopy in adult rats adapted to altered gravity. AQP1 expression was decreased by 64% at the apical pole of choroidal cells in rats dissected 5.5-8 h after a 14-day spaceflight. AQP1 was significantly overexpressed in rats readapted for 2 days to Earth's gravity after an 11-day flight (48% overshoot, when compared with the value measured in control rats). In a ground-based model that simulates some effects of weightlessness and alters choroidal structures and functions, apical AQP1 expression was reduced by 44% in choroid plexus from rats suspended head down for 14 days and by 69% in rats suspended for 28 days. Apical AQP1 was rapidly enhanced in choroid plexus of rats dissected 6 h after a 14-day suspension (57% overshoot, in comparison with control rats) and restored to the control level when rats were dissected 2 days after the end of a 14-day suspension. Decreases in the apical expression of choroidal AQP1 were also noted in rats adapted to hypergravity in the NASA 24-ft centrifuge: AQP1 expression was reduced by 47% and 85% in rats adapted for 14 days to 2 G and 3 G, respectively. AQP1 is downregulated in the apical membrane of choroidal cells in response to altered gravity and is rapidly restored after readaptation to normal gravity. This suggests that water transport, which is partly involved in the choroidal production of cerebrospinal fluid, might be decreased during spaceflight and after chronic hypergravity.

Animals↗
PubMed2002

Effect of gravity on chest wall mechanics.

Chest wall mechanics was studied in four subjects on changing gravity in the craniocaudal direction (G(z)) during parabolic flights. The thorax appears very compliant at 0 G(z): its recoil changes only from -2 to 2 cmH(2)O in the volume range of 30-70% vital capacity (VC). Increasing G(z) from 0 to 1 and 1.8 G(z) progressively shifted the volume-pressure curve of the chest wall to the left and also caused a fivefold exponential decrease in compliance. For lung volume <30% VC, gravity has an inspiratory effect, but this effect is much larger going from 0 to 1 G(z) than from 1 to 1.8 G(z). For a volume from 30 to 70% VC, the effect is inspiratory going from 0 to 1 G(z) but expiratory from 1 to 1.8 G(z). For a volume greater than approximately 70% VC, gravity always has an expiratory effect. The data suggest that the chest wall does not behave as a linear system when exposed to changing gravity, as the effect depends on both chest wall volume and magnitude of G(z).

Compliance↗
PubMed2006

Modeling gravity-dependent plasticity of the angular vestibuloocular reflex with a physiologically based neural network.

A neural network model was developed to explain the gravity-dependent properties of gain adaptation of the angular vestibuloocular reflex (aVOR). Gain changes are maximal at the head orientation where the gain is adapted and decrease as the head is tilted away from that position and can be described by the sum of gravity-independent and gravity-dependent components. The adaptation process was modeled by modifying the weights and bias values of a three-dimensional physiologically based neural network of canal-otolith-convergent neurons that drive the aVOR. Model parameters were trained using experimental vertical aVOR gain values. The learning rule aimed to reduce the error between eye velocities obtained from experimental gain values and model output in the position of adaptation. Although the model was trained only at specific head positions, the model predicted the experimental data at all head positions in three dimensions. Altering the relative learning rates of the weights and bias improved the model-data fits. Model predictions in three dimensions compared favorably with those of a double-sinusoid function, which is a fit that minimized the mean square error at every head position and served as the standard by which we compared the model predictions. The model supports the hypothesis that gravity-dependent adaptation of the aVOR is realized in three dimensions by a direct otolith input to canal-otolith neurons, whose canal sensitivities are adapted by the visual-vestibular mismatch. The adaptation is tuned by how the weights from otolith input to the canal-otolith-convergent neurons are adapted for a given head orientation.

Adaptation, Physiological↗