Hydrogen sulfide inhibits Cav3.2 T-type Ca2+ channels

The importance of H2S as a physiological signaling molecule continues to develop, and ion channels are emerging as a major family of target proteins through which H2S exerts many actions. The purpose of the present study was to investigate its effects on T-type Ca2+ channels. Using patch-clamp electrophysiology, we demonstrate that the H2S donor, NaHS (10 μM-1 mM) selectively inhibits Cav3.2 T-type channels heterologously expressed in HEK293 cells, whereas Cav3.1 and Cav3.3 channels were unaffected. The sensitivity of Cav3.2 channels to H2S required the presence of the redox-sensitive extracellular residue H191, which is also required for tonic binding of Zn2+ to this channel. Chelation of Zn2+ with N,N,N',N'-tetra-2-picolylethylenediamine prevented channel inhibition by H2S and also reversed H2S inhibition when applied after H2S exposure, suggesting that H2S may act via increasing the affinity of the channel for extracellular Zn2+ binding. Inhibition of native T-type channels in 3 cell lines correlated with expression of Cav3.2 and not Cav3.1 channels. Notably, H2S also inhibited native T-type (primarily Cav3.2) channels in sensory dorsal root ganglion neurons. Our data demonstrate a novel target for H2S regulation, the T-type Ca2+ channel Cav3.2, and suggest that such modulation cannot account for the pronociceptive effects of this gasotransmitter.

Oxygen-dependent hydroxylation by Factor Inhibiting HIF (FIH) regulates the TRPV3 ion channel

Factor Inhibiting HIF (FIH) is an oxygen-dependent asparaginyl hydroxylase that regulates the hypoxia-inducible factors (HIFs). Several proteins containing ankyrin repeat domains have been characterised as substrates of FIH, although there is little evidence for a functional consequence of hydroxylation on these substrates. This study demonstrates that the transient receptor potential vanilloid 3 (TRPV3) channel is hydroxylated by FIH on asparagine 242 within the cytoplasmic ankyrin repeat domain. Hypoxia, FIH inhibitors and mutation of asparagine 242 all potentiated TRPV3-mediated current, without altering TRPV3 protein levels, indicating that oxygen-dependent hydroxylation inhibits TRPV3 activity. This novel mechanism of channel regulation by oxygendependent asparaginyl hydroxylation is likely to extend to other ion channels.

Modeling the observed proton aurora and ionospheric convection responses to changes in the IMF clock angle: 2. Persistence of ionospheric convection

We apply a numerical model of time-dependent ionospheric convection to two directly driven reconnection pulses during a 15-min interval of southward IMF on 26 November 2000. The model requires an input magnetopause reconnection rate variation, which is here derived from the observed variation in the upstream IMF clock angle, q. The reconnection rate is mapped to an ionospheric merging gap, the MLT extent of which is inferred from the Doppler-shifted Lyman-a emission on newly opened field lines, as observed by the FUV instrument on the IMAGE spacecraft. The model is used to reproduce a variety of features observed during this event: SuperDARN observations of the ionospheric convection pattern and transpolar voltage; FUV observations of the growth of patches of newly opened flux; FUVand in situ observations of the location of the Open-Closed field line Boundary (OCB) and a cusp ion step. We adopt a clock angle dependence of the magnetopause reconnection electric field, mapped to the ionosphere, of the form Enosin4(q/2) and estimate the peak value, Eno, by matching observed and modeled variations of both the latitude, LOCB, of the dayside OCB (as inferred from the equatorward edge of cusp proton emissions seen by FUV) and the transpolar voltage FPC (as derived using the mapped potential technique from SuperDARN HF radar data). This analysis also yields the time constant tOCB with which the open-closed boundary relaxes back toward its equilibrium configuration. For the case studied here, we find tOCB = 9.7 ± 1.3 min, consistent with previous inferences from the observed response of ionospheric flow to southward turnings of the IMF. The analysis confirms quantitatively the concepts of ionospheric flow excitation on which the model is based and explains some otherwise anomalous features of the cusp precipitation morphology.

An analysis of the accuracy of magnetopause reconnection rate variations deduced from cusp ion dispersion characteristics

We present an analysis of the accuracy of the method introduced by Lockwood et al. (1994) for the determination of the magnetopause reconnection rate from the dispersion of precipitating ions in the ionospheric cusp region. Tests are made by applying the method to synthesised data. The simulated cusp ion precipitation data are produced by an analytic model of the evolution of newly-opened field lines, along which magnetosheath ions are firstly injected across the magnetopause and then dispersed as they propagate into the ionosphere. The rate at which these newly opened field lines are generated by reconnection can be varied. The derived reconnection rate estimates are then compared with the input variation to the model and the accuracy of the method assessed. Results are presented for steady-state reconnection, for continuous reconnection showing a sine-wave variation in rate and for reconnection which only occurs in square wave pulses. It is found that the method always yields the total flux reconnected (per unit length of the open-closed field-line boundary) to within an accuracy of better than 5%, but that pulses tend to be smoothed so that the peak reconnection rate within the pulse is underestimated and the pulse length is overestimated. This smoothing is reduced if the separation between energy channels of the instrument is reduced; however this also acts to increase the experimental uncertainty in the estimates, an effect which can be countered by improving the time resolution of the observations. The limited time resolution of the data is shown to set a minimum reconnection rate below which the method gives spurious short-period oscillations about the true value. Various examples of reconnection rate variations derived from cusp observations are discussed in the light of this analysis.

Location and characteristics of the reconnection X line deduced from low-altitude satellite and ground-based observations: 2. Defense Meteorological Satellite Program and European Incoherent Scatter data

We present an analysis of a cusp ion step, observed by the Defense Meteorological Satellite Program (DMSP) F10 spacecraft, between two poleward moving events of enhanced ionospheric electron temperature, observed by the European Incoherent Scatter (EISCAT) radar. From the ions detected by the satellite, the variation of the reconnection rate is computed for assumed distances along the open-closed field line separatrix from the satellite to the X line, do. Comparison with the onset times of the associated ionospheric events allows this distance to be estimated, but with an uncertainty due to the determination of the low-energy cutoff of the ion velocity distribution function, ƒ(ν). Nevertheless, the reconnection site is shown to be on the dayside magnetopause, consistent with the reconnection model of the cusp during southward interplanetary magnetic field (IMF). Analysis of the time series of distribution function at constant energies, ƒ(ts), shows that the best estimate of the distance do is 14.5±2 RE. This is consistent with various magnetopause observations of the signatures of reconnection for southward IMF. The ion precipitation is used to reconstruct the field-parallel part of the Cowley D ion distribution function injected into the open low-latitude boundary layer in the vicinity of the X line. From this reconstruction, the field-aligned component of the magnetosheath flow is found to be only −55±65 km s−1 near the X line, which means either that the reconnection X line is near the stagnation region at the nose of the magnetosphere, or that it is closely aligned with the magnetosheath flow streamline which is orthogonal to the magnetosheath field, or both. In addition, the sheath Alfvén speed at the X line is found to be 220±45 km s−1, and the speed with which newly opened field lines are ejected from the X line is 165±30 km s−1. We show that the inferred magnetic field, plasma density, and temperature of the sheath near the X line are consistent with a near-subsolar reconnection site and confirm that the magnetosheath field makes a large angle (>58°) with the X line.

EISCAT observations of ion composition and temperature anisotropy in the high-latitude F-region

The papers by Winser et al. [(1990) J. atmos. terr. Phys.52, 501] and Häggström and Collis [(1990) J. atmos. terr. Phys.52, 519] used plasma flows and ion temperatures, as measured by the EISCAT tristatic incoherent scatter radar, to investigate changes in the ion composition of the ionospheric F-layer at high latitudes, in response to increases in the speed of plasma convection. These studies reported that the ion composition rapidly changed from mainly O+ to almost completely (>90%) molecular ions, following rapid increases in ion drift speed by >1 km s−1. These changes appeared inconsisent with theoretical considerations of the ion chemistry, which could not account for the large fractions of molecular ions inferred from the obsevations. In this paper, we discuss two causes of this discrepancy. First, we reevaluate the theoretical calculations for chemical equilibrium and show that, if we correct the derived temperatures for the effect of the molecular ions, and if we employ more realistic dependences of the reaction rates on the ion temperature, the composition changes derived for the faster convection speeds can be explained. For the Winser et al. observations with the radar beam at an aspect angle of ϕ = 54.7° to the geomagnetic field, we now compute a change to 89% molecular ions in < 2 min, in response to the 3 km s−1 drift. This is broadly consistent with the observations. But for the two cases considered by Häggström and Collis, looking along the field line (ϕ = 0°), we compute the proportion of molecular ions to be only 4 and 16% for the observed plasma drifts of 1.2 and 1.6 km s−1, respectively. These computed proportions are much smaller than those derived experimentally (70 and 90%). We attribute the differences to the effects of non-Maxwellian, anisotropic ion velocity distribution functions. We also discuss the effect of ion composition changes on the various radar observations that report anisotropies of ion temperature.

DE-2 observations of filamentary currents at ionospheric altitudes

Conjunctive measurements made by the Dynamics Explorer 1 and 2 spacecraft on October 22, 1981, under conditions of southward IMF, suggest the existence of a cusp ion injection from a region at the magnetopause with a scale size of ∼ 1/2 to 1 R E . Current signatures observed by the LAPI and MAGB instruments on board DE-2 indicate the existence of a rotation in the magnetic field that is consistent with a filamentary current system. The observed current structure can be interpreted as the ionospheric signature of a flux transfer event (FTE). In addition to this large-scale current structure there exist three small-scale filamentary current pairs. These current pairs close locally and thus, if our interpretation of this event as an FTE is correct, represent the first reported observations of FTE interior structure at low-altitudes.

Incoherent scatter radar observations of non-Maxwellian ion velocity distributions in the auroral F-region

Observations by the EISCAT experiments “POLAR” and Common Programme CP-3 reveal non-Maxwellian ion velocity distributions in the auroral F-region ionosphere. Analysis of data from three periods is presented. During the first period, convection velocities are large (≈2 km s-1) and constant over part of a CP-3 latitude scan; the second period is one of POLAR data containing a short-lived (<1 min.) burst of rapid (>1.5 km s-1) flow. We concentrate on these two periods as they allow the study of a great many features of the ion-neutral interactions which drive the plasma non-thermal and provide the best available experimental test for models of the 3-dimensional ion velocity distribution function. The third period is included to illustrate the fact that non-thermal plasma frequently exists in the auroral ionosphere: the data, also from the POLAR experiment, cover a three-hour period of typical auroral zone flow and analysis reveals that the ion distribution varies from Maxwellian to the threshold of a toroidal form.

Ion flows and heating at a contracting polar-cap boundary

Data recorded by the POLAR experiment run on the EISCAT radar during the international GISMOS campaign of 3–5 June 1987 are studied in detail. The polar-cap boundary, as denned by an almost shear East-West convection reversal, was observed to jump southward across the EISCAT field of view in two steps at 02:00 and 03:00 Magnetic Local Time and subsequently to contract back between 04:00 and 07:00 M.L.T. An annulus of enhanced ion temperature and non-thermal plasma was observed immediately equatorward of the contracting boundary due to the lag in the response of the neutral-wind pattern to the change in ion flows. The ion flow at the boundary is shown to be relatively smooth at 15 s resolution and directed northward, with velocities which exceed that of the boundary itself. The effect of velocity shears on the beamswinging technique used to derive the ion flows is analyzed in detail and it is shown that, for certain orientations of the cap boundary, spurious flows into the cap can be generated. However, these are much smaller than the observed flows into the polar cap and cannot explain the potential difference across the observed segment of the cap boundary (extending over 2 h of M.L.T.) which is roughly 7 kV. Similarly, an observed slowing of the zonal flow near the boundary cannot be explained as an error introduced by the use of the beamswinging technique. The results could be interpreted as being due to reconnection occurring on the dawn flank of the magnetopause (mapping to the polar cap at 04:30 06:30 M.L.T.). However, they are more consistent with recent observations of slow anti-sunward flow of closed field lines on the flanks of the geomagnetic tail, which appears to be generated by some form of “viscous” coupling to the magnetosheath plasma.

Vertical ion flow in the polar ionosphere

Outflowing ions from the polar ionosphere fall into two categories: the classical polar wind and the suprathermal ion flows. The flows in both these categories vary a great deal with altitude. The classical polar wind is supersonic at high altitude: at ∼3 RE geocentric, the observed polar wind is H+ dominated and has a Mach number of 2.5–5.1. At 400–600 km, thermal and suprathermal upward O+ ion fluxes frequently occur at the poleward edge of the nightside auroral oval during magnetically active times. Above 500 km, ions are accelerated transverse to the local geomagnetic field. At 1400 km, transversely accelerated ions are frequently observed in winter nights but rarely appear in the summer. In the dayside cleft above ∼2000 km, ions of all species are transversely heated and upwell with significant number and heat fluxes, forming a cleft ion fountain as they convect across the polar cap. Upwelling ions are observed most (least) frequently in the summer (winter). At yet higher altitudes, energetic (>10 eV to several kiloelectron volts) upflowing H+ and O+ ions are frequently observed, their active time occurrence frequency being as high as 0.7 at auroral latitudes and 0.3 in the polar cap. Their composition, intensity, and angular characteristics vary quantitatively with solar activity, being O+ dominant and more intense near solar maximum. Their resulting ion outflow is dominated by ions below 1 keV and reaches 3.5×10^26 O+ and 7×10^25 H+ ions s^{−1} at magnetically active times (Kp≥5) near solar maximum. In comparison, the estimated polar wind ion outflow at times of moderate solar activity is 7×10^25H+ and 4×10^24 He+ ions s^{−1}. The estimated <10-eV cleft ion fountain flow is 3.8×10^25 O+ and 8.6×10^23 H+ ions s^{−1} near solar maximum.

The modelled occurrence of non-thermal plasma in the ionospheric F-region and the possible consequences for ion outflows into the magnetosphere

A global, time-dependent, three-dimensional, coupled ionosphere-thermosphere model is used to predict the spatial distribution of non-thermal plasma in the F-layer. It is shown that, even for steady-state conditions with Kp as low as 3, the difference between the ion and neutral velocities often exceeds the neutral thermal speed by a factor, D', which can be as large as 4. Theoretically, highly non-Maxwellian, and probably toroidal, ion velocity distributions are expected when D' exceeds about 1.5. The lack of response of the neutral winds to sunward ion drifts in the dawn sector of the auroral oval cause this to be the region most likely to contain toroidal distributions. The maximum in D' is found in the throat region of the convection pattern, where the strong neutral winds of the afternoon sector meet the eastward ion flows of the morning sector. These predictions are of interest, not only to radar scientists searching for non-thermal ionospheric plasma, but also as one possible explanation of the initial heating and upward flows of ions in the cleft ion fountain and nightside auroral oval, both of which are a major source of plasma for the magnetosphere.

Non-Maxwellian ion velocity distributions observed using EISCAT

Recent observations from the EISCAT incoherent scatter radar have revealed bursts of poleward ion flow in the dayside auroral ionosphere which are consistent with the ionospheric signature of flux transfer events at the magnetopause. These bursts frequently contain ion drifts which exceed the neutral thermal speed and, because the neutral thermospheric wind is incapable of responding sufficiently rapidly, toroidal, non-Maxwellian ion velocity distributions are expected. The EISCAT observations are made with high time resolution (15 seconds) and at a large angle to the geomagnetic field (73.5°), allowing the non-Maxwellian nature of the distribution to be observed remotely for the first time. The observed features are also strongly suggestive of a toroidal distribution: characteristic spectral shape, increased scattered power (both consistent with reduced Landau damping and enhanced electric field fluctuations) and excessively high line-of-sight ion temperatures deduced if a Maxwellian distribution is assumed. These remote sensing observations allow the evolution of the distributions to be observed. They are found to be non-Maxwellian whenever the ion drift exceeds the neutral thermal speed, indicating that such distributions can exist over the time scale of the flow burst events (several minutes).

Low-energy ion outflows from the ionosphere during a major polar cap expansion — evidence for equatorward motion of inverted-V structures

Data from the Dynamics Explorer 1 satellite and the EISCAT and Sondrestrom incoherent scatter radars, have allowed a study of low-energy ion outflows from the ionosphere into the magnetosphere during a rapid expansion of the polar cap. From the combined radar data, a 200kV increase in cross-cap potential is estimated. The upflowing ions show “X” signatures in the pitch angle-time spectrograms in the expanding midnight sector of the auroral oval. These signatures reveal low-energy (below about 60eV), light-ion beams sandwiched between two regions of ion conics and are associated with inverted-V electron precipitation. The lack of mass dispersion of the poleward edge of the event, despite great differences in the times of flight, reflects the equatorward expansion of the acceleration regions at velocities similar to those of the antisunward convection. In addition, a transient burst of upflow of 0+ is observed within the cap, possibly due to enhanced Joule heating during the event.

Low-energy ion flows into the magnetosphere

Ion flows from the ionosphere into the magnetosphere fall into two main categories: cold (<1eV), “classical” polar wind and heated (>1eV), suprathermal ion outflows. A wealth of new understanding of these outflows has resulted from the Dynamics Explorer Mission. This review describes both the confirmation of the predicted classical polar wind as well as the revelation of a great variety of low-energy suprathermal outflows: the cleft ion fountain, the nightside auroral fountaion (X-events, toroids and field-aligned flows) and polar cap outflows. The main emphasis is placed on flows at energies below about 50eV, observed by the Retarding Ion Mass Spectrometer (RIMS) on board the Dynamics Explorer 1 satellite; limited comparisons are made with results from other instruments which sample different energy ranges.

Upwelling O+ ion source characteristics

Recent observations from the Dynamics Explorer 1 (DE-1) spacecraft have shown that the dayside auroral zone is an important source of very low-energy superthermal O^+ ions for the polar magnetosphere. When observed at 2000- to 5000-km altitude, the core of the O^+ distribution exhibits transverse heating to energies on the order of 10 eV, significant upward heat flux, and subsonic upward flow at significant flux levels exceeding 10^8 cm^{-2}s^{-1}. The term "upwelling ions" has been adopted to label these flows, which stand out in sharp contrast to the light ion polar wind flows observed in the same altitude range in the polar cap and subauroral magnetosphere. We have chosen a typical upwelling ion event for detailed study, correlating retarding ion mass spectrometer observations of the low-energy plasma with energetic ion observations and local electromagnetic field observations. The upwelling ion signature is colocated with the magnetospheric cleft as marked by precipitating energetic magnetosheath ions. The apparent ionospheric heating is clearly linked with the magnetic field signatures of strong field-aligned currents in the vicinity of the dayside polar cap boundary. Electric field and ion plasma measurements indicate that a very strong and localized convection channel or jet exists coincident with the other signatures of this event. These observations indicate that transverse ion heating to temperatures on the order of 10^5 K in the 2000- to 5000-km ionosphere is an important factor in producing heavy ion outflows into the polar magnetosphere. This result contrasts with recent suggestions that electron heating to temperatures of order 10^4 K is the most important parameter with regard to O^+ outflow.