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M A Shea

Publications and source records attributed to M A Shea.

At least 37 records · Page 2Linked to original sources

The heliolongitudinal distribution of solar flares associated with solar proton events.

We find that the heliolongitudinal distribution of solar flares associated with earth-observed solar proton events is a function of the particle measurement energy. For solar proton events containing fluxes with energies exceeding 1 GeV, we find a Gaussian distribution about the probable root of the Archimedean spiral favorable propagation path leading from the earth to the sun. This distribution is modified as the detection threshold is lowered. For > 100 MeV solar proton events with fluxes > or = 10 protons (cm2-sec-ster)-1 we find the distribution becomes wider with a secondary peak near the solar central meridian. When the threshold is lowered to 10 MeV the distribution further evolves. For > 10 MeV solar proton events having a flux threshold at 10 protons (cm2-sec-ster)-1 the distribution can be considered to be a composite of two Gaussians. One distribution is centered about the probable root of the Archimedean spiral favorable propagation path leading from the earth to the sun, and the other is centered about the solar central meridian. For large flux solar proton events, those with flux threshold of 1000 (cm2-sec-ster)-1 at energies > 10 MeV, we find the distribution is rather flat for about 40 degrees either side of central meridian.

Earth, Planet↗

Effects of orbit progression on the radiation exposures from solar proton fluxes in low Earth orbit under geomagnetic storm conditions.

The present study examines the effects of orbit progression on the exposures within a Space Station Freedom module in a 51.6-degree inclined orbit at 450 km. The storm evolution is modeled after the November 1960 event, and the solar proton flux evolution is taken from the August 1972 solar proton event. The effects of a strong magnetic shock, such as was observed during the October 1989 event, is also modeled. The statistics on hourly average storm fields for the last forty years reveal that the largest geomagnetic storms approach a Dst value of -500 nanotesla at the storm peak. Similarly, one of the largest satellite-measured proton flux (> 10 MeV) for space exposures is the event of August 1972. The effects of orbit progression (advance of the line of nodes) is examined for the above conditions to study the variation of exposures under differing times of occurrence of the solar proton peak intensity, attainment of geomagnetic storm maximum, and the location of the line of nodes of the last geomagnetically protected orbit. The impact of the inherent inhomogeneity of the space station module is examined as a limiting factor on exposure with regard to the need of additional parasitic shielding.

Algorithms↗

Solar-interplanetary circumstances associated with the major events in March and June 1991 and comparison with similar events of previous solar cycles.

The solar activity and geomagnetic storm events of March and June 1991 were associated with the appearance of an enhanced particle flux in the trapped radiation belts as discovered by the CRRES satellite observations and later measured by shuttle radiation detectors. The solar-interplanetary conditions associated with these events appear to be a major sequence of activity near the sun's central meridian generating powerful fast interplanetary shocks resulting in major perturbations to the magnetosphere. The solar-interplanetary events in 1991 are discussed and compared to similar activity in the past such as the events in February 1986, August 1972, July 1961, November 1960, and July 1959.

Earth, Planet↗

Calcium binding decreases the stokes radius of calmodulin and mutants R74A, R90A, and R90G.

Calmodulin (CaM) is an intracellular cooperative calcium-binding protein essential for activating many diverse target proteins. Biophysical studies of the calcium-induced conformational changes of CaM disagree on the structure of the linker between domains and possible orientations of the domains. Molecular dynamics studies have predicted that Ca4(2+)CaM is in equilibrium between an extended and compact conformation and that Arg74 and Arg90 are critical to the compaction process. In this study gel permeation chromatography was used to resolve calcium-induced changes in the hydrated shape of CaM at pH 7.4 and 5.6. Results showed that mutation of Arg 74 to Ala increases the R(s) as predicted; however, the average separation of domains in Ca4(2+)-CaM was larger than predicted by molecular dynamics. Mutation of Arg90 to Ala or Gly affected the dimensions of apo-CaM more than those of Ca4(2+)-CaM. Calcium binding to CaM and mutants (R74A-CaM, R90A-CaM, and R90G-CaM) lowered the Stokes radius (R(s)). Differences between R(s) values reported here and Rg values determined by small-angle x-ray scattering studies illustrate the importance of using multiple techniques to explore the solution properties of a flexible protein such as CaM.

Amino Acid Sequence↗

Discontinuous equilibrium titrations of cooperative calcium binding to calmodulin monitored by 1-D 1H-nuclear magnetic resonance spectroscopy.

Calmodulin binds up to four calcium ions cooperatively in response to cellular signaling events. To understand the functional energetics of calcium activation of calmodulin, it is important to monitor individual Ca(2+)-binding sites and other positions at partial degrees of saturation. This study is the first use of 1-D proton NMR to monitor the equilibrium Ca(2+)-binding properties of calmodulin. Protein concentrations required for NMR experiments (approximately 1 mM) are approximately 1000-fold greater than the Kd values for calcium binding to calmodulin, preventing a direct continuous equilibrium titration of calmodulin. Thus, dialysates of calmodulin in buffers of experimentally determined [Ca2+]free were prepared to conduct discontinuous equilibrium titrations at both 92 and 152 mM KCl. For the C-terminal domain, the normalized area of the delta-protons of Y138 defined calcium binding isotherms. For N-terminal domain resonances (F16C delta H, T26C alpha H, D64C alpha H, and F65C delta H), the calcium-dependent change in chemical shift defined isotherms. These are the first residue-specific studies to monitor the energetics of Ca2+ binding to the N-terminal domain in wild-type holo calmodulin. Calcium binding to both domains appeared cooperative and binding affinity decreased in higher KCl. Isotherms resolved from the side chain resonances of F16 and F65 had a lower median ligand activity and a slightly higher degree of cooperativity than isotherms resolved from the backbone resonances of D64 and T26. Salt-dependent changes in apparent intradomain cooperativity differed for the domains: at higher salt, delta Gc increased for the C-terminal domain while remaining constant or decreasing for the N-terminal domain.

Amino Acid Sequence↗

Quantitative endoproteinase GluC footprinting of cooperative Ca2+ binding to calmodulin: proteolytic susceptibility of E31 and E87 indicates interdomain interactions.

Calmodulin is the primary eukaryotic intracellular calcium receptor. Cooperative calcium binding to two sites in each of two domains drives large conformational changes that enable it to activate target proteins. An understanding of the molecular mechanism of cooperativity requires determination of the conformational states populated by calmodulin, the intrinsic free energies of binding calcium to four sites, and the nature and degree of intradomain and interdomain interactions. To monitor residue-specific conformational changes within calmodulin as calcium binds, we have developed a new quantitative proteolytic footprinting method using endoproteinase GluC (EndoGluC). Under conditions of very limited proteolysis, cleavage occurred at only five of the sixteen positions possible in calmodulin. The relative abundance of fragments indicated that calcium induced changes in the susceptibility of individual peptide bonds. Quantitative susceptibility profiles were resolved for two positions: E31-L32, in site I in the N-terminal domain, and E87-A88, preceding site III in the C-terminal domain. In apocalmodulin, E87-A88 was susceptible to EndoGluC; calcium binding to sites III and IV caused monotonic protection from proteolysis. The response of E31-L32 was biphasic. In apocalmodulin, it was resistant to cleavage. Susceptibility was induced by calcium binding to sites III and IV, followed by protection induced by calcium binding to sites I and II. This indicated that calmodulin must adopt at least three distinguishable conformations and suggested that the two domains interact. Model-dependent equilibrium constants were resolved from the EndoGluC susceptibility profiles for E31 and E87; they indicated cooperative binding within each domain. Approaches taken to validate this proteolytic footprinting method are described.

Amino Acid Sequence↗

Bohr effects of the partially-ligated (CN-met) intermediates of hemoglobin as probed by quaternary assembly.

Free energies of quaternary assembly (dimers to tetramers) were determined for the 10 ligation species of CN-methemoglobin in the region of the alkaline Bohr effect (pH 7.0-9.5). Analysis of this database yielded the following principal findings: (1) At each pH, the nine CN-met species exhibit two distinct values of Bohr proton release and Bohr free energy. The two Bohr effects are found to distribute in a fashion that coincides with predictions of a symmetry rule (Ackers et al., 1992), i.e., the first value reflects a "tertiary Bohr effect" arising from ligation within the quaternary T tetramer and a second Bohr effect arises from the quaternary transition (T-->R) which occurs when both dimeric half-molecules acquire at least one ligated subunit. (2) The Bohr effects for CN-met ligation are in good agreement with previously-established Bohr effects for stepwise O2 binding under identical conditions (Chu et al., 1984). (3) In combination with recent studies which show that CN-met species [21] has a quaternary T structure (Daugherty et al., 1991; Doyle & Ackers, 1992; LiCata et al., 1993), the present results show that the "tertiary Bohr effect" within quaternary T exceeds the Bohr effect of dissociated dimers, as suggested by Lee and Karplus (1983). (4) The tertiary Bohr effect is found to account for the pH dependence of tertiary constraint energy, delta Gtc, which "pays" for ligand-binding cooperativity prior to the quaternary (T-->R) switchover. Possible origins of the tertiary Bohr effect and its relationship to the quaternary Bohr effect are considered.

Allosteric Regulation↗

Significant proton events of solar cycle 22 and a comparison with events of previous solar cycles.

The sun has produced several high energy and large fluence solar proton events during solar cycle 22. This recent activity is similar to activity that occurred in the 19th solar cycle before the advent of routine space measurements. In a review of the recent events and a comparison with significant solar proton events of previous solar cycles, it appears that the 20th and 21st solar cycles were deficient in the total fluence of solar particles as detected at the earth. Therefore, when models of maximum solar proton fluxes to be encountered for deep space missions are developed, solar proton data acquired during the present solar cycle should be incorporated.

Astronomy↗

The dose rate observed on 19-21 October 1989 and its modulation by geophysical effects.

The Liulin dosimeter-radiometer on the MIR space station detected the 19 October 1989 high energy solar proton event. These results show that the main particle increase contains protons with energies up to about 9 GeV. After the main particle onset the Liulin dosimeter observed a typical geomagnetic cutoff modulation of the dose rate from the solar particles as the MIR space station traversed magnetic latitudes. When the interplanetary shock and associated solar plasma enveloped the earth on 20 October between 14 and 17 UT the radiation exposure increased significantly due to the lowering of the geomagnetic cutoff. The analysis of this event shows how various geophysical phenomena can significantly modulate the dose rate encountered by earth-orbiting spacecraft.

Cosmic Radiation↗

Proton flare and magnetic storm effect in the vicinity of the Earth.

We have developed a model and associated computational procedure for estimating energetic proton exposures during a major solar proton event that occur in combination with a large magnetic storm. Transmission functions for solar protons are computed using geomagnetic vertical cutoff data for quiescent and disturbed conditions. Predicted exposures in low altitude polar orbit are found to be orders of magnitude greater for severe magnetic storm conditions than are corresponding exposures in the absence of major disturbances. We examine the response scenario for the events of November 1960 as an example.

Computer Simulation↗

Geomagnetic cutoffs: a review for space dosimetry applications.

The earth's magnetic field acts as a shield against charged particle radiation from interplanetary space, technically described as the geomagnetic cutoff. The cutoff rigidity problem (except for the dipole special case) has "no solution in closed form". The dipole case yields the Stormer equation which has been repeatedly applied to the earth in hopes of providing useful approximations of cutoff rigidities. Unfortunately the earth's magnetic field has significant deviations from dipole geometry, and the Stormer cutoffs are not adequate for most applications. By application of massive digital computer power it is possible to determine realistic geomagnetic cutoffs derived from high order simulation of the geomagnetic field. Using this technique, "world-grids" of directional cutoffs for the earth's surface and for a limited number of satellite altitudes have been derived. However, this approach is so expensive and time consuming it is impractical for most spacecraft orbits, and approximations must be used. The world grids of cutoff rigidities are extensively used as lookup tables, normalization points and interpolation aids to estimate the effective geomagnetic cutoff rigidity of a specific location in space. We review the various options for estimating the cutoff rigidity for earth-orbiting satellites.

Cosmic Radiation↗

Modeling the time-intensity profile of solar flare generated particle fluxes in the inner heliosphere.

It is possible to model the time-intensity profile of solar particles expected in space after the occurrence of a significant solar flare on the sun. After the particles are accelerated in the flare process, if conditions are favorable, they may be released into the solar corona and then into space. The heliolongitudinal gradients observed in the inner heliosphere are extremely variable, reflecting the major magnetic structures in the solar corona which extend into space. These magnetic structures control the particle gradients in the inner heliosphere. The most extensive solar particle measurements are those observed by earth-orbiting spacecraft, and forecast and prediction procedures are best for the position of the earth. There is no consensus of how to extend the earth-based models to other locations in space. Local interplanetary conditions and structures exert considerable influence on the time-intensity profiles observed. The interplanetary shock may either reduce or enhance the particle intensity observed at a specific point in space and the observed effects are very dependent on energy.

Astronomical Phenomena↗

Identification of the intermediate allosteric species in human hemoglobin reveals a molecular code for cooperative switching.

The 10 ligation species of human cyanomethemoglobin were previously found to distribute into three discrete cooperative free energy levels according to a combinatorial code (i.e., dependent on both the number and configuration of ligated subunits). Analysis of this distribution showed that the hemoglobin tetramer occupies a third allosteric state in addition to those of the unligated (T) and fully ligated (R) species. To determine the nature of the intermediate allosteric state, we have studied the effects of pH, temperature, and single-site mutations on its free energy of quaternary assembly, in parallel with corresponding data on the deoxy (T) and fully ligated (R) species. Results indicate that the intermediate allosteric tetramer has the deoxy (T) quaternary structure. This finding, together with the resolved energetic distribution of the 10 microstates reveals a symmetry rule for quaternary switching--i.e., switching from T to R occurs whenever a binding step creates a tetramer with one or more ligated subunits on each side of the alpha 1 beta 2 intersubunit contact. These studies also reveal significant cooperativity within each alpha 1 beta 1 dimer of the T-state tetramer. The ligand-induced tertiary free energy alters binding affinity within the T structure by 170-fold prior to quaternary switching.

Allosteric Site↗

Subunit hybridization studies of partially ligated cyanomethemoglobins using a cryogenic method. Evidence for three allosteric states.

Reaction of tetrameric hemoglobin with ligands at the four heme sites yields nine species that have structurally unique combinations of ligated and unligated subunits. Using hemoglobins where the ligated subunits contain cyanomethemoglobin, Smith and Ackers studied the dimer-tetramer assembly reactions in all nine of the partially ligated species (F. R. Smith and G. K. Ackers, Proc. Natl. Acad. Sci. U.S.A. 82 (1985) 5347). They found a third assembly free energy in addition to those of unligated hemoglobin and fully ligated cyanomethemoglobin. The observed distribution of the three assembly free energies among the ten species was found to be incompatible with the two-state mechanism of allosteric control (J. Monod, J. Wyman and J. P. Changeaux, J. Mol. Biol. 12 (1965) 81). The results indicated a mechanism of 'combinatorial switching' in which the binding free energies per site change with configuration of occupied sites and not just their number. In this study, we have confirmed the existence of three assembly free energies among the ten ligation species using a cryogenic method (M. Perrella and L. Rossi-Bernardi, Methods Enzymol. 76 (1981) 133). For one of the species we find a different free energy assignment from that reported by Smith and Ackers; for all other species we observe the same assignments as in earlier work. The revised distribution also requires a 'combinatorial' mechanism of allosteric switching among the three states.

Freezing↗

PPS-87: a new event oriented solar proton prediction model.

A new event-oriented solar proton prediction model has been developed and implemented at the USAF Space Environment forecast facility. This new model generates predicted solar proton time-intensity profiles for a number of user adjustable energy ranges and is also capable of making predictions for the heavy ion flux. The computer program is designed so a forecaster can select inputs based on the data available in near real-time at the forecast center as the solar flare is occurring. The predicted event amplitude is based on the electromagnetic emission parameters of the solar flare (either microwave or soft X-ray emission) and the solar flare position on the sun. The model also has an update capability where the forecaster can normalize the prediction to actual spacecraft observations of spectral slope and particle flux as the event is occurring in order to more accurately predict the future time-intensity profile of the solar particle flux. Besides containing improvements in the accuracy of the predicted energetic particle event onset time and magnitude, the new model converts the predicted solar particle flux into an expected radiation dose that might be experienced by an astronaut during EVA activities or inside the space shuttle.

Computer Simulation↗