Subcellular components of the amphibian egg: insights provided by gravitational studies.

Most cytoplasmic regions of fertilized amphibian eggs move with respect to the gravity vector in experimentally gravity oriented eggs. The pattern and extent of this movement varies among different batches of eggs. This variation in apparent cytoplasmic viscosity (or, conversely, cytoplasmic mobility) can be correlated with variations in subsequent morphogenesis of experimental, gravitationally manipulated eggs. Therefore, the proper interpretation of gravity experiments with amphibian eggs requires that one understand the subcellular basis for this variation on cytoplasmic mobility. Variation in the packing of the major cytoplasmic organelle, the yolk platelets, or the organization and amount of cytoskeletal components may explain the variation in cytoplasmic mobility. Evidence is presented that the variation in yolk volume density (fraction of total cytoplasmic volume occupied by yolk platelets) does not account for the variation in cytoplasmic mobility in Xenopus laevis eggs. Experimental evidence from cold-shocked inverted eggs indicates that microtubules may be involved in determining cytoplasmic mobility. However, quantitative evidence that the microtubule levels and state of the microtubules (polymerized vs. non-polymerized) in the whole Xenopus laevis egg does not correlate directly with cytoplasmic mobility is presented. The apparent conflict these data represent regarding the role of the cytoskeleton in determining cytoplasmic mobility is discussed.     

Amphibian egg cytoplasm response to altered g-forces and gravity orientation

Elucidation of dorsal/ventral polarity and primary embryonic axis development in amphibian embryos requires an understanding of cytoplasmic rearrangements in fertile eggs at the biophysical, physiological, and biochemical levels. Evidence is presented that amphibian egg cytoplasmic components are compartmentalized. The effects of altered orientation to the gravitational vector (i.e., egg inversion) and alterations in gravity force ranging from hypergravity (centrifugation) to simulated microgravity (i.e., horizontal clinostat rotation) on cytoplasmic compartment rearrangements are reviewed. The behavior of yolk compartments as well as a newly defined (with monoclonal antibody) non-yolk cytoplasmic compartment, in inverted eggs and in eggs rotated on horizontal clinostats at their buoyant density, is discussed.     

Fertilization of sea urchin eggs in space and subsequent development under normal conditions.

Sea urchin eggs are generally considered as most suitable animal models for studying fertilization processes and embryonic development. In the present study, they are used for determining a possible role of gravity in fertilization and the establishment of egg polarity and the embryonic axis. For this purpose, eggs of the particularly well known and suitable species Paracentrotus lividus have been automatically fertilized under microgravity conditions during the Swedish sounding rocket flights MASER IV and MASER V. It turns out, that fertilization "in Space" occurs normally and that subsequent embryonic and larval development of such eggs, continued on the ground, is normal, leading to advanced pluteus stages.     

Paramecium--a model system for studying cellular graviperception.

Experiments under varied gravitational accelerations as well as in density-adjusted media showed that sensation of gravity in protists may be linked to the known principles of mechanosensation. Paramecium, a ciliate with clear graviresponses (gravitaxis and gravikinesis) is an ideal model system to prove this hypothesis since the ciliary activity and thus the swimming behaviour is controlled by the membrane potential. It has also been assumed that the cytoplasmic mass causes a distinct stimulation of the bipolarly distributed mechano-sensitive K+ and Ca2+ ion channels in the plasma membrane in dependence of the spatial orientation of the cell. In order to prove this hypothesis, different channel blockers are currently under investigation. Gadolinium did not inhibit gravitaxis in Paramecium, showing that it does not specifically block gravireceptors. Further studies concentrated on the question of whether second messengers are involved in the gravity signal transduction chain. Exposure to 5 g for up to 10 min led to a significant increase in cAMP.     

Regional gravity field model in Pakistan area from the combination of CHAMP,GRACE and ground data using least squares collocation: A case study

This study describes a methodology of recovery of the Earth’s gravity field from CHAMP and GRACE satellites data in Pakistan using least squares collocation (LSC) based downward continuation technique. The CHAMP height anomalies and GRACE gravity disturbances derived from the observed satellite data have been used in combination solution using LSC with observed gravity values at the Earth surface. The combined covariance functions of height anomalies and/or gravity disturbances at satellite altitudes and observed gravity anomalies at Earth surface have been used as the basis for combination and downward continuation solution. The variance of predicted gravity anomalies from GRACE gravity disturbances is relatively lower than the corresponding results of gravity anomalies from CHAMP height anomalies. This fact may be attributed partly to the amplification of noise and partly to the unstable inverse transformation process of height anomalies to gravity anomalies. The impact of data error variance has been studied in the context of smoothing and noise reduction in the final solution of downward continuation using least squares collocation. The raising of data error suppresses the noise and as a result a smooth final solution is obtained. The prediction results appear to be dependent on the quality of data and goodness of combined covariance function, which are fairly comparable for the CHAMP and GRACE data. The recovered gravity field from satellite data appears to contribute mainly to medium and long wavelength parts of total gravity field spectrum. Due to flexibility of data handling in least squares collocation, this procedure is applicable to any observable of gravity field being at different altitudes and with different data spacing.     

AstroNewt: early development of newt in space.

AstroNewt experiment explores the effects of earth gravity on the early development of Japanese red-bellied newt, Cynops pyrrhogaster. Since female newts keep spermatophore in cloaca, fertilized eggs could be obtained without mating. Fertilization of newt's egg occurs just prior to spawning, so that gonadotrophic cues applied to females in orbit leads to laying eggs fertilized just in space. A property of newt being kept in hibernation at low temperature may be of great help for the space experiment carried out with much limited resources. A general outline of the AstroNewt project is shown here in addition to some technical advances for the development of the project. Experimental schemes of two space experiments (IML-2 in summer 1994 and unmanned SFU at the beginning of 1995) are also shown.     

Cytoskeleton and gravity at work in the establishment of dorso-ventral polarity in the egg of Xenopus laevis

The establishment of polarities during early embryogenesis is essential for normal development. Amphibian eggs are appropriate models for studies on embryonic pattern formation. The animal-vegetal axis of the axially symmetrical amphibian egg originates during oogenesis and foreshadows the main body axis of the embryo. The dorso-ventral polarity is epigenetically established before first cleavage. Recent experiments strongly suggest that in the monospermic eggs of the anuran Xenopus laevis both the cytoskeleton and gravity act in the determination of the dorso-ventral polarity. In order to test the role of gravity in this process, eggs will be fertilized under microgravity conditions during the SL-D1 flight in 1985. In a fully automatic experiment container eggs will be kept under well-defined conditions and artificially fertilized as soon as microgravity is reached; eggs and embryos at different stages will then be fixed for later examination. Back on earth the material will be analysed and we will know whether fertilization under microgravity conditions is possible. If so, the relation of the dorso-ventral axis to the former sperm entry point will be determined on the whole embryos; in addition eggs and embryos will be analysed cytologically.     

The temporal variation of mean gravity of atmosphere

One of the precise widely used global Zenith Hydrostatic Delay (ZHD) model is based on the gravity value at the centroid of the atmospheric column at the station of observation and gravity value at the centroid is constant in this model for a specific location throughout the year. However, as the content and extent of atmosphere varies temporally, its centroid and consequently gravity value at the centroid also varies. Apart from this, the actual atmospheric condition of different region is not alike. Therefore, there is a need to develop a regional mean gravity model and development of such model has been discussed in this paper. To obtain the mean gravity model, first a regional model of centroid height of atmospheric column was developed as a function of the surface pressure and temperature. It was developed by multiple regressions between estimated centroid of the atmosphere and surface pressure, surface temperature using radiosonde data of five radiosonde stations spread over the Indian subcontinent. Three years radiosonde data from 2006 to 2008 was used for each station. The root mean square error in estimating centroid of the atmospheric column is about ±326 m, which is negligible considering the variability of the atmosphere and its content. The centroid height model has been used to formulate the mean gravity model, considering uniform lapse rate in gravity with height. It is found that proposed mean gravity model provides temporal variation of mean gravity values at the centroid and thus matches with the reality. The interesting advantage of the developed model is that the model shows diurnal variation of mean gravity. The accuracy of ZHD has shown of the order of about 0.3 mm using the developed regional mean gravity model. However, already developed ZHD model has shown a slight inferior result compared to the developed model. These models have shown accuracy of about 0.8 mm and 0.6 mm.     

Seasonal variation of total electron content at crest of equatorial anomaly station during low solar activity conditions

The variability of total electron content (TEC) over the crest of equatorial anomaly station Bhopal has been studied during the low solar activity period (2005–2006) using global positioning system (GPS) data. Diurnal variation of TEC is studied for different seasons. Interesting features like the winter anomaly, semiannual anomaly and noon bite out in TEC have been reported. GPS derived TEC is then compared with International Reference Ionosphere (IRI) 2001 model and the difference between predictions and observation is being studied. Using the variability index we have also studied the TEC variability for different seasons and also during quiet and disturbed conditions. A higher variability is observed on quiet days as compared to disturbed days during daytime and nighttime hours.     

Simultaneous bi-station measurements of mesospheric waves from Indian low latitudes

Coordinated mesospheric night-airglow measurements have been carried out from two stations, Gadanki (13.5° N, 79.2° E) and Allahabad (25.5° N, 81.9° E), India during April 2009 to study the common gravity wave features. With two nights of coordinated measurements we find some of the wave periodicities to be similar at the two locations. Simultaneous OH and O₂ intensity measurements over Gadanki reveal these features to be upward propagating gravity waves while the coordinated OH intensity measurements of similar waves from Allahabad show the large spatial extent of these waves.     

A direct approach to the study of the effect of gravity on axis formation in birds.

A system has been developed to enable the normal development of aborted very early uterine avian embryos, outside the female's uterus. The shell-less aborted egg was put into a foster shell of a sister egg, previously laid by the same female. The empty space between the shell and aborted egg was filled with artificial uterine fluid. The reconstructed eggs were incubated at 42 degrees C for 30 hours in a vertical position. The atmosphere contained a high concentration of CO2 (8-10%). At the termination of the 30 h the eggs were transferred to incubation at 37 degrees C in normal atmospheric conditions. Normal development has been recorded for a certain percentage of eggs incubated up to 12 days. In other cases abnormalities, arrested development or development of extraembryonic membranes only, without a sign of an embryonic axis, have been observed. The three important conclusions from the above experiments were: 1. It is possible to develop a closed, self-contained system, disconnected from the female's body, that would support the development of early uterine embryos. 2. The incidence of embryo-less extraembryonic membranes in such a system, is correlated with the degree of detachment of the "yolk" from the outer envelopes. 3. Such a system can be further developed into an experiment suited for microgravity conditions which will be an alternative to an experiment with live birds. The experiment will be aimed at testing the importance of gravity in changing the radially symmetrical avian blastoderm into a bilaterally symmetrical blastoderm.     

GOCE orbit analysis: Long-wavelength gravity field determination using the acceleration approach

The restricted sensitivity of the Gravity field and steady-state Ocean Circulation Explorer (GOCE) gradiometer instrument requires satellite gravity gradiometry to be supplemented by orbit analysis in order to resolve long-wavelength features of the geopotential. For the hitherto published releases of the GOCE time-wise (TIM) and GOCE space-wise gravity field series—two of the official ESA products—the energy conservation method has been adopted to exploit GPS-based satellite-to-satellite tracking information. On the other hand, gravity field recovery from data collected by the CHAllenging Mini-satellite Payload (CHAMP) satellite showed the energy conservation principle to be a sub-optimal choice. For this reason, we propose to estimate the low-frequency part of the gravity field by the point-wise solution of Newton’s equation of motion, also known as the acceleration approach. This approach balances the gravitational vector with satellite accelerations, and hence is characterized by (second-order) numerical differentiation of the kinematic orbit. In order to apply the method to GOCE, we present tailored processing strategies with regard to low-pass filtering, variance–covariance information handling, and robust parameter estimation. By comparison of our GIWF solutions (initials GI for “Geodätisches Institut” and IWF for “Institut für WeltraumForschung”) and the GOCE-TIM estimates with a state-of-the-art gravity field solution derived from GRACE (Gravity Recovery And Climate Experiment), we conclude that the acceleration approach is better suited for GOCE-only gravity field determination as opposed to the energy conservation method.     

Calcium signaling in plant cells in altered gravity.

Changes in the intracellular Ca2+ concentration in altered gravity (microgravity and clinostating) evidence that Ca2+ signaling can play a fundamental role in biological effects of microgravity. Calcium as a second messenger is known to play a crucial role in stimulus-response coupling for many plant cellular signaling pathways. Its messenger functions are realized by transient changes in the cytosolic ion concentration induced by a variety of internal and external stimuli such as light, hormones, temperature, anoxia, salinity, and gravity. Although the first data on the changes in the calcium balance in plant cells under the influence of altered gravity have appeared in 80th, a review highlighting the performed research and the possible significance of such Ca2+ changes in the structural and metabolic rearrangements of plant cells in altered gravity is still lacking. In this paper, an attempt was made to summarize the available experimental results and to consider some hypotheses in this field of research. It is proposed to distinguish between cell gravisensing and cell graviperception; the former is related to cell structure and metabolism stability in the gravitational field and their changes in microgravity (cells not specialized to gravity perception), the latter is related to active use of a gravitational stimulus by cells presumebly specialized to gravity perception for realization of normal space orientation, growth, and vital activity (gravitropism, gravitaxis) in plants. The main experimental data concerning both redistribution of free Ca2+ ions in plant cell organelles and the cell wall, and an increase in the intracellular Ca2+ concentration under the influence of altered gravity are presented. Based on the gravitational decompensation hypothesis, the consequence of events occurring in gravisensing cells not specialized to gravity perception under altered gravity are considered in the following order: changes in the cytoplasmic membrane surface tension --> alterations in the physicochemical properties of the membrane --> changes in membrane permeability, --> ion transport, membrane-bound enzyme activity, etc. --> metabolism rearrangements --> physiological responses. An analysis of data available on biological effects of altered gravity at the cellular level allows one to conclude that microgravity environment appears to affect cytoskeleton, carbohydrate and lipid metabolism, cell wall biogenesis via changes in enzyme activity and protein expression, with involvement of regulatory Ca2+ messenger system. Changes in Ca2+ influx/efflux and possible pathways of Ca2+ signaling in plant cell biochemical regulation in altered gravity are discussed.     

Investigation of the characteristics of wavelike oscillations of post-sunset equatorial ionospheric irregularity by decomposing fluctuating TEC

Post-sunset ionospheric irregularities are common features of the equatorial ionosphere that affect radio communication and navigation systems; their triggering physical mechanism is not yet fully understood. Atmospheric gravity wave is considered as a seeding mechanism for the occurrence of ionospheric irregularities (Abdu et al., 2009). To understand the effects of atmospheric waves, characteristics of wavelike oscillation from ionospheric total electron content (TEC) fluctuation that can be obtained from superposition of different oscillation modes have been investigated. Decomposing fluctuating TEC into different oscillation modes and investigating oscillation characteristics of each component is also important to get insight about the characteristics of individual atmospheric waves that may cause TEC fluctuation. In this paper we have investigated characteristics of components of fluctuating TEC obtained from SCINDA GPS receiver installed at Bahir Dar, (geographic coordinate, 11.5°N, 37.6° E, and dip latitude of 2.5°N) Ethiopia during April 2012. First Empirical Mode Decomposition (EMD) has been applied to decompose TEC fluctuation into different oscillation modes that are known as Intrinsic Mode Function (IMF). Hilbert-Huang Transform (HHT) and Continuous Wavelet Transform (CWT) have been applied to investigate the characteristics of wave-like oscillations. Applying EMD on fluctuating vTEC corresponding to a GPS satellite, five components are found. Results from HHT and CWT have shown excellent agreement. In addition, it is found out that the median periods of oscillation of those five components are 9, 17, 47, 78, and 118 min. Of these periods, 17 and 47 min respectively are oscillation periods of components of TEC fluctuation with occurrence frequency of 92% and 91% that may be interpreted as the manifestation of two frequently occurring components of atmospheric gravity waves that are likely generated by the motion of solar terminator.     

Regulative development of Xenopus laevis in microgravity.

To test whether gravity is required for normal amphibian development, Xenopus laevis females were induced to ovulate aboard the orbiting Space Shuttle. Eggs were fertilized in vitro, and although early embryonic stages showed some abnormalities, the embryos were able to regulate and produce nearly normal larvae. These results demonstrate for the first time that a vertebrate can ovulate in the virtual absence of gravity, and that the eggs can develop to a free-living stage.     

Long-wavelength lunar gravity field recovery from simulated orbit and inter-satellite tracking data

As has been demonstrated recently, inter-satellite Ka-band tracking data collected by the GRAIL (Gravity Recovery And Interior Laboratory) spacecraft have the potential to improve the resolution and accuracy of the lunar gravity field by several orders of magnitude compared to previous models. By means of a series of simulation studies, here we investigate the contribution of inter-satellite ranging for the recovery of the Moon’s gravitational features; the evaluation of results is made against findings from ground-based Doppler tracking. For this purpose we make use of classical dynamic orbit determination, supported by the analysis of satellite-to-satellite tracking observations. This study sheds particularly light on the influence of the angular distance between the two satellites, solar radiation modeling and the co-estimation of the lunar Love number k₂. The quality of the obtained results is assessed by gravity field power spectra, gravity anomalies and precision orbit determination. We expect our simulation results to be supportive for the processing of real GRAIL data.     

Effect of gravity field on the nonequilibrium/nonlinear chemical oscillation reactions.

Biological systems have evolved for a long time under the normal gravity. The Belousov-Zhabotinsky (BZ) reaction is a nonlinear chemical system far from the equilibrium that may be considered as a simplified chemical model of the biological systems so as to study the effect of gravity. The reaction solution is comprised of bromate in sulfuric acid as an oxidizing agent, 1,4-cyclohexanedione as an organic substrate, and ferroin as a metal catalyst. Chemical waves in the BZ reaction-diffusion system are visualized as blue and red patterns of ferriin and ferroin, respectively. After an improvement to the tubular reaction vessels in the experimental setup, the traveling velocity of chemical waves in aqueous solutions was measured in time series under normal gravity, microgravity, hyper-gravity, and normal gravity using the free-fall facility of JAMIC (Japan Microgravity Center), Hokkaido, Japan. Chemical patterns were collected as image data via CCD camera and analyzed by the software of NIH image after digitization. The estimated traveling velocity increased with increasing gravity as expected. It was clear experimentally that the traveling velocity of target patterns in reaction diffusion system was influenced by the effect of convection and correlated closely with the gravity field.     

An improved altimeter-derived gravity anomaly from shipborne gravity based on the mean sea surface height constraint factors method

Coastal marine gravity modeling faces challenges due to the degradation of the quality and poor coverage of altimeter data in coastal regions. The effective fusion of shipborne gravity data and altimeter-derived marine gravity data can make shipborne gravity data more useful for the accurate estimation of altimeter-derived coastal marine gravity. A mean sea surface height constraint factor (MSSHCF) method based on the ordinary kriging method and the remove-restore technique is proposed to fuse altimeter-derived gravity model with shipborne gravity data. In this method, all data are standardized during the interpolation process to reduce the error and mean sea surface as a vertical variable is added to the semi-variance function in ordinary kriging to obtain the residual shipborne gravity as corrected data source. The coastal marine gravity models V2.1 and V3.1 which fused altimeter-derived gravity data with shipborne gravity data and V1.1 without shipborne gravity data at a spatial resolution of 1′×1′ can be obtained. Validation experiments show that the accuracy of the gravity model V3.1 obtained by the MSSHCF method more closely agrees with the validated gravity model DTU17 and SS V31 than the model V2.1 obtained by the ordinary kriging interpolation method and the V1.1 model. Our results were validated against shipborne gravity data; the accuracy of model V3.1 was 4.95 % higher than the model V1.1 in South China Sea area A and 2.48 % higher in South China Sea area B. Meanwhile, the accuracy of model V3.1 was 2.07 % higher than model V2.1 in South China Sea area A and 2.42 % higher in South China Sea area B. The effects of distance from the coast and sea depth on the marine gravity model were also evaluated. The results show that the gravity model V3.1 has higher accuracy with the change in ocean distance and depth than the V2.1 and V1.1 gravity models. Thus, our study shows that the MSSHCF method effectively refines coastal altimeter-derived gravity using shipborne gravity data.     

Remote sounding of atmospheric gravity waves with satellite limb and nadir techniques

Recent advances in satellite techniques hold great potential for mapping global gravity wave (GW) processes at various altitudes. Poor understanding of small-scale GWs has been a major limitation to numerical climate and weather models for making reliable forecasts. Observations of short-scale features have important implication for validating and improving future high-resolution numerical models. This paper summarizes recent GW observations and sensitivities from several satellite instruments, including MLS, AMSU-A, AIRS, GPS, and CLAES. It is shown in an example that mountain waves with horizontal wavelengths as short as 30 km now can be observed by AIRS, reflecting the superior horizontal resolution in these modern satellite instruments. Our studies show that MLS, AMSU-A and AIRS observations reveal similar GW characteristics, with the observed variances correlated well with background winds. As a complementary technique, limb sounding instruments like CRISTA, CLAES, and GPS can detect GWs with better vertical but poorer horizontal resolutions. To resolve different parts of the broad GW spectrum, both satellite limb and nadir observing techniques are needed, and a better understanding of GW complexities requires joint analyses of these data and dedicated high-resolution model simulations.     

Evaluation of experiments involving the study of plant orientation and growth under different gravitational conditions.

The manifestation of gravitropic reaction in plants has been considered from the phylogenetic point of view. A chart has been suggested according to which it is supposed that the first indications of the ability to identify the direction of the gravitational vector were inherent in the most ancient eukaryotes, which gave rise to green, brown, yellow-green, golden and diatomaceous algae as well as fungi. The experiments on the role of gravity in plant ontogenesis are being continued. The sum total of the data obtained in a number of experiments in space shows that under these conditions a structurally modified but normally functioning gravireceptive apparatus is formed. The data confirming the modification, under changed gravity, of the processes of integral and cellullar growth of the axial organs of seedlings as well as of the anatomo-morphological structure and developmental rates of plants during their prolonged growth in space are presented. It is assumed that this fact testifies to the presence of systems interacting with gravity during plant ontogenesis. At the same time the necessity for further experiments in order to differentiate an immediate biological effect of gravity from the ones conditioned by it indirectly due to the changes in the behavior of liquids and gases is pointed out. The methodological aspects of biological experiments in space as the main source of reliable information on the biological role of gravity are discussed.