Particle Ejection Contributions to the Rotational Acceleration and Orbit Evolution of Asteroid (101955) Bennu.

This paper explores the implications of the observed Bennu particle ejection events for that asteroid's spin rate and orbit evolution, which could complicate interpretation of the Yarkovsky-O'Keefe-Radzievskii-Paddack (YORP) and Yarkovsky effects on this body's spin rate and orbital evolution. Based on current estimates of particle ejection rates, we find that the overall contribution to Bennu's spin and orbital drift is small or negligible as compared to the Yarkovsky and YORP effects. However, if there is a large unseen component of smaller mass ejections or a strong directionality in the ejection events, it could constitute a significant contribution that could mask the overall YORP effect. This means that the YORP effect may be stronger than currently assumed. The analysis is generalized so that the particle ejection effect can be assessed for other bodies that may be subject to similar mass loss events. Further, our model can be modified to address different potential mechanisms of particle ejection, which are a topic of ongoing study.

Heterogeneous mass distribution of the rubble-pile asteroid (101955) Bennu.

The gravity field of a small body provides insight into its internal mass distribution. We used two approaches to measure the gravity field of the rubble-pile asteroid (101955) Bennu: (i) tracking and modeling the spacecraft in orbit about the asteroid and (ii) tracking and modeling pebble-sized particles naturally ejected from Bennu's surface into sustained orbits. These approaches yield statistically consistent results up to degree and order 3, with the particle-based field being statistically significant up to degree and order 9. Comparisons with a constant-density shape model show that Bennu has a heterogeneous mass distribution. These deviations can be modeled with lower densities at Bennu's equatorial bulge and center. The lower-density equator is consistent with recent migration and redistribution of material. The lower-density center is consistent with a past period of rapid rotation, either from a previous Yarkovsky-O'Keefe-Radzievskii-Paddack cycle or arising during Bennu's accretion following the disruption of its parent body.

Hayabusa2 arrives at the carbonaceous asteroid 162173 Ryugu-A spinning top-shaped rubble pile.

The Hayabusa2 spacecraft arrived at the near-Earth carbonaceous asteroid 162173 Ryugu in 2018. We present Hayabusa2 observations of Ryugu's shape, mass, and geomorphology. Ryugu has an oblate "spinning top" shape, with a prominent circular equatorial ridge. Its bulk density, 1.19 ± 0.02 grams per cubic centimeter, indicates a high-porosity (>50%) interior. Large surface boulders suggest a rubble-pile structure. Surface slope analysis shows Ryugu's shape may have been produced from having once spun at twice the current rate. Coupled with the observed global material homogeneity, this suggests that Ryugu was reshaped by centrifugally induced deformation during a period of rapid rotation. From these remote-sensing investigations, we identified a suitable sample collection site on the equatorial ridge.

The dynamic geophysical environment of (101955) Bennu based on OSIRIS-REx measurements.

The top-shape morphology of asteroid (101955) Bennu is commonly found among fast-spinning asteroids and binary asteroid primaries, and might have contributed significantly to binary asteroid formation. Yet a detailed geophysical analysis of this morphology for a fast-spinning asteroid has not been possible prior to the Origins, Spectral Interpretation, Resource Identification, and Security-Regolith Explorer (OSIRIS-REx) mission. Combining the measured Bennu mass and shape obtained during the Preliminary Survey phase of OSIRIS-REx, we find a significant transition in Bennu's surface slopes within its rotational Roche lobe, defined as the region where material is energetically trapped to the surface. As the intersection of the rotational Roche lobe with Bennu's surface has been most recently migrating towards its equator (given Bennu's increasing spin rate), we infer that Bennu's surface slopes have been changing across its surface within the last million years. We also find evidence for substantial density heterogeneity within this body, suggesting that its interior has a distribution of voids and boulders. The presence of such heterogeneity and Bennu's top-shape is consistent with spin-induced failure at some point in its past, although the manner of its failure cannot be determined yet. Future measurements by the OSIRIS-REx spacecraft will give additional insights and may resolve questions regarding the formation and evolution of Bennu's top-shape morphology and its link to the formation of binary asteroids.

The efficiency of sensory information coding by mechanoreceptor neurons.

Most sensory systems encode external signals into action potentials for transmission to the central nervous system, but little is known about the cost or efficiency of this encoding. We measured the information capacity at three stages of encoding in the neurons of a spider slit-sense mechanoreceptor organ. For the receptor current under voltage clamp, the capacity was approximately 1400 bits/s, but when the neuron was allowed to generate a receptor potential, nonlinear membrane processes improved the capacity to >2000 bits/s. Finally, when action potentials were produced, the capacity dropped to approximately 200 bits/s, or approximately 14% of the receptor current capacity. These measurements provide a quantitative estimation of the cost of encoding analog signals into action potentials.

Information processing by graded-potential transmission through tonically active synapses.

Many neurons use graded membrane-potential changes, instead of action potentials, to transmit information. Traditional synaptic models feature discontinuous transmitter release by presynaptic action potentials, but this is not true for synapses between graded-potential neurons. In addition to graded and continuous transmitter release, they have multiple active zones, ribbon formations and L-type Ca2+ channels. These differences are probably linked to the high rate of vesicle fusion required for continuous transmitter release. Early stages of sensory systems provide some of the best characterized graded-potential neurons, and recent work on these systems suggests that modification of synaptic transmission by adaptation is a powerful feature of graded synapses.

Evidence that channels below 1 pS cause the volume-sensitive chloride conductance in T84 cells.

The volume-activated chloride current of T84 human colonic cells was studied using the whole-cell patch clamp. The current appeared reliably with a mild osmotic gradient and in the absence of intracellular ATP. It reversed at the chloride equilibrium potential and was blocked by the chloride channel blocker DIDS. Development of the current was accompanied by an increase in the current noise variance, typical of increasing ion channel open probability. Noise variance was always well-fitted by a double Lorentzian relationship with corner frequencies at approximately 1.7 Hz and approximately 60 Hz. The increase in variance during development of the volume-sensitive current was mostly due to an increase in the high frequency component. The relationship between noise variance and membrane current was well-fitted by a relationship with a single channel conductance of approximately 0.2 pS.

A large conductance, Ca2+-activated K+ channel in a human lung epithelial cell line (A549).

A large conductance, Ca2+-activated K+ channel in a human lung epithelial cell line (A549) was identified using the single channel patch clamp technique. Channel conductance was 242 +/- 33 pS (n = 67) in symmetrical KCl (140 mM). The channel was activated by membrane depolarization and increased cytosolic Ca2+. High selectivity was observed for K+ over Rb+(0.49) > Cs+(0.14) > Na+(0.09). Open probability was significantly decreased by Ba2+ (5 mM) and quinidine (5 mM) to either surface, but TEA (5 mM) was only effective when added to the external surface. All effects were reversible. Increasing cytosolic Ca2+ concentration from 10(-7) to 10(-6) M caused an increase in open probability from near zero to fully activated. ATP decreased open probability at approximately 2 mM, but the effect was variable. The channel was almost always observed together with a smaller conductance channel, although they could both be seen individually. We conclude that A549 cells contain large conductance Ca2+-activated K+ channels which could explain a major fraction of the K+ conductance in human alveolar epithelial membranes.

Halide permeation through 10 pS and 20 pS anion channels in human airway epithelial cells.

Halide permeability sequences were obtained from reversal potential measurements of single-channel currents through 10 pS and 20 pS anion channels in human airway epithelial cells. The sequences obtained were Cl- greater than I- greater than Br- greater than or equal to F- for the 10 pS channel and Cl- greater than I- greater than or equal to Br- greater than or equal to F- for the 20 pS channel. However, the permeability differences were not large, the greatest being 0.66 for the ratio of fluoride to chloride permeability in the 20 pS channel. Single-channel currents were also measured with solutions of constant halide concentration but varying ratios of chloride to fluoride ions. An anomalous mole fraction effect was observed for the 20 pS channel but not for the 10 pS channel, suggesting that the former is a multi-ion channel. Comparison of the halide permeability sequences of these two channels with those of whole-cell currents in other epithelial cells does not support their involvement in any of the known whole-cell epithelial currents.

Subconductance states in calcium-activated potassium channels from canine airway smooth muscle.

The single-channel patch clamp technique was used to analyze subconductance states in the 260 pS calcium-activated potassium channel from canine airway smooth muscle. More than sixty minutes of single channel data (greater than 87,000 events) from five excised patches were analyzed. Six subconductance amplitudes were clearly established to be 17, 33, 41, 52, 63 and 72% of the full conductance. Subconductance openings were usually brief (milliseconds) and represented less than 5% of the total channel open time, but they also persisted for several seconds on rare occasions. They appeared to be unaffected by voltage or time after seal formation, but may have increased in occurrence with decreasing calcium concentration. Irregular amplitude intervals, and the presence of ramp-like, analog transitions between conductance states, suggest a model for maxi-K subconductance states in which the channel protein undergoes random conformational changes causing a variable pore size.

Rapid coating of glass-capillary microelectrodes for single-electrode voltage-clamp.

The single-electrode voltage-clamp technique requires sharp glass-capillary microelectrodes, whose electrical properties often limit the capabilities of the recording system. Here, we describe a rapid and simple way of coating fine microelectrodes with Dricote and Vaseline that improves their performance during voltage-clamp. The coating prevented clogging of the tips, improved the capacitance compensation of the electrodes, helped to seal the electrode tips into cell membranes and allowed visualization of the tips under saline solution. This new coating method led to greatly improved recordings and better characterization of the transduction and voltage-activated currents in an isolated preparation of spider mechanosensory neurons.

Ouabain selectively affects the slow component of sensory adaptation in an insect mechanoreceptor.

Sensory activity in the cockroach tactile spine neuron adapts rapidly to a step deflection. This rapid adaptation is caused by a rise in the threshold for action potential production, which has two components with different time constants and drug sensitivities. The basis of the slow component is unknown but it is insensitive to a wide range of ion channel blockers. In the present experiments the slow component was selectively reduced by ouabain, suggesting that it is due to the activation of an electrogenic sodium pump.

Mapping extracellular excitability in an insect mechanoreceptor neuron.

The cockroach tactile spine contains a single bipolar mechanosensory neuron. Extracellular stimulation of the neuron is possible by cutting the spine and lowering a microelectrode into the lumen, where the neuron is located, but neither the microelectrode nor the neuron can be visualized during stimulation. The threshold for electrical stimulation of the neuron was measured as a function of spatial position in the lumen. The spine was then fixed and serially sectioned for computer-aided reconstruction. Alignment of threshold measurements with reconstructions produced maps of excitability around the neuron. The lowest threshold was always close to the sensory dendrite or the adjacent soma. These results are discussed in terms of models of action potential initiation in this class of sensory neurons.

Estimated single-channel conductance of mechanically-activated channels in a spider mechanoreceptor.

Noise analysis was used to estimate the single-channel conductance and number of channels responsible for the mechanically-activated current in the sensory neurons of a spider mechanoreceptor organ. External slits of the VS-3 slit-sense organ in the patellar cuticle of Cupiennius salei were moved with a piezoelectric stimulator while glass microelectrodes penetrated the adjacent cell bodies. Receptor currents were measured by the switching single-electrode voltage clamp technique during both step and ramp displacements of the slits. Current records were segmented in time, and the variance and amplitude of the current were obtained from each segment, to allow fitting of the variance vs. amplitude relationship by a standard equation based on a two-state channel. Mean values of 7.5 pS and 253 were obtained for the conductance and number of channels from 75 separate recordings. These values are in good agreement with the small number of other estimates of these parameters from different mechanoreceptor preparations.

Dissociation and culture of mechanosensory neurons for patch clamp analysis.

Single mechanosensory neurons were isolated from chordotonal organs of adult cockroach antennae. Transmission and scanning electron microscopy showed that the soma and part of the mechanosensory ending survived the dissociation. In culture, outgrowth occurred from the ending. Regions of cell membrane were accessible for patch clamp analysis and channels were recorded. The ability to record channel activity in isolated mechanosensory neurons will allow the study of mechanotransduction mechanisms at the membrane level.

Octopamine selectively modifies the slow component of sensory adaptation in an insect mechanoreceptor.

The effects of octopamine were studied on the dynamic behavior of the sensory neuron in the cockroach femoral tactile spine. The neuron is a rapidly adapting mechanoreceptor in which adaptation occurs by elevation of the threshold for action potential encoding. The threshold follows increases or decreases of membrane potential, with a delay that involves two separate exponential components. Previous evidence has associated the slow component with sodium pumping and the fast component with sodium channel inactivation. Octopamine reversibly raised the resting threshold and increased but slowed the slow component. These data indicate that octopamine has specific effects on membrane-ionic processes in insect sensory neurons.

Postsynaptic dorsal column and cuneate neurons in raccoon: comparison of response properties and cross-correlation analysis.

The responses of 111 postsynaptic dorsal column (PSDC) neurons in the cervical spinal cord and 51 cuneate neurons with receptive fields on the glabrous skin of the forepaw were studied in anesthetized raccoons using extracellular recording techniques. The PSDC neurons had larger receptive fields than the cuneate neurons, but in both groups the fields never extended onto hairy skin. PSDC and cuneate neurons had approximately the same mean latency to electrical stimulation of the receptive field, but PSDC neurons had significantly lower thresholds. The majority of both PSDC and cuneate neurons also responded to electrical stimulation of an adjacent digit, even though they did not respond to mechanical stimulation of that digit. Cross-correlation analysis of the activity of 51 pairs of PSDC and cuneate neurons recorded simultaneously revealed a significant interaction in 26 pairs during spontaneous activity. In 20 of these neuron pairs, the probability that the cuneate neuron would fire was greater after the PSDC neuron had fired (suggesting a spinocuneate interaction), five pairs showed an interaction in the opposite (cuneospinal) direction, and one pair had a significant inhibitory interaction. These interactions occurred more often when the receptive fields of the two neurons were overlapping than when their fields were on adjacent digits. Frequency response analysis revealed greater coherence for those pairs showing a spinocuneate interaction than for those with a cuneospinal interaction. These results support the hypothesis that the PSDC system exerts a tonic facilitatory effect on cuneate neurons and that there may be some somatotopic organization to the interactions. However, the similar response latencies of the two groups of neurons makes it unlikely that PSDC neurons could contribute to the rapid initial processing of cutaneous information by the cuneate nucleus.

Sodium channel distribution in a spider mechanosensory organ.

A site-directed antibody was used immunocytochemically to measure the distribution of sodium channels in the tissues of a spider mechanoreceptor organ. The VS-3 slit sense organ contains 7-8 pairs of bipolar sensory neurons; these neurons are representative of a wide range of arthropod mechanoreceptors. Sensory transduction is thought to occur at the tips of the dendrites and to cause action potentials that are regeneratively conducted to the cell bodies, although it has not been possible to confirm this by direct intracellular recordings from the dendrites. Wholemount preparations were labelled by immunofluorescence and thin sections were immunogold labelled, using an antibody to the highly conserved SP19 sequence of the voltage-activated sodium channel. Labelling for sodium channels was found in the neurons and in their surrounding glial cells. Both cytoplasm and membranes were labelled, but immunogold particles were clearly aligned along cell membranes, indicating that the majority of labelling represented membrane-bound sodium channels. Channel density in the dendrites was similar to the axons and higher than in the cell bodies, supporting the idea of active conduction in the sensory dendrites. Labelling in glial cell membranes was indistinguishable from the neighboring neurons, suggesting a significant role for sodium channels in the functions of these supporting cells.

The time course of sensory adaptation in the cockroach tactile spine.

Power-law dynamics have been widely used to fit the adaptation of sensory receptors, including the cockroach tactile spine. We used a new log-binning technique to re-examine step responses in the tactile spine. The power-law only fitted responses over a restricted time period, while a sum of three exponential decays gave an accurate fit over the entire response duration.

Information transmission at 500 bits/s by action potentials in a mechanosensory neuron of the cockroach.

Action potentials are widely used to transmit information within nervous systems but information encoding and transmission rates by action potentials are poorly understood. In the absence of knowledge about encoding, most previous work has used signal-to-noise ratios to estimate information capacities. We used a mechanosensory neuron to transmit information by a simple encoding scheme that allowed us to measure the transmission rate directly. Using either mechanical or electrical stimulation, information was transmitted at rates up to 500 bits/s, higher than ever reported before for real action potentials. However, the maximum possible message length decreased strongly with transmission rate, from approximately infinite at 100 bits/s to approximately 100 ms at 500 bits/ s, probably due to ionic adaptation processes within the neuronal membrane.