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.

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.

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.

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.

Frequency response functions and information capacities of paired spider mechanoreceptor neurons.

Pseudorandom white-noise stimulation followed by direct spectral estimation was used to obtain linear frequency response and coherence functions from paired, but dynamically different, spider mechanosensory neurons. The dynamic properties of the two neuron types were similar with either mechanical or electrical stimulation, showing that action potential encoding dominates the dynamics. Phase-lag data indicated that action potential initiation occurs more rapidly during mechanical stimulation, probably in the distal sensory dendrites. Total information capacity, calculated from coherence, as well as information per action potential, were both similar in the two types of neurons, and similar to the few available estimates from other spiking neurons. However, information capacity and information per action potential both depended strongly on neuronal firing rate, which has not been reported before.

Inactivation of voltage-activated Na(+) currents contributes to different adaptation properties of paired mechanosensory neurons.

Voltage-activated sodium current (I(Na)) is primarily responsible for the leading edge of the action potential in many neurons. While I(Na) generally activates rapidly when a neuron is depolarized, its inactivation properties differ significantly between different neurons and even within one neuron, where I(Na) often has slowly and rapidly inactivating components. I(Na) inactivation has been suggested to regulate action potential firing frequency in some cells, but no clear picture of this relationship has emerged. We studied I(Na) in both members of the paired mechanosensory neurons of a spider slit-sense organ, where one neuron adapts rapidly (type A) and the other slowly (type B) in response to a step depolarization. In both neuron types I(Na) activated and inactivated with single time constants of 2--3 ms and 5--10 ms, respectively, varying with the stimulus intensity. However, there was a clear difference in the steady-state inactivation properties of the two neuron types, with the half-maximal inactivation (V(50)) being -40.1 mV in type A neurons and -58.1 mV in type B neurons. Therefore I(Na) inactivated closer to the resting potential in the more slowly adapting neurons. I(Na) also recovered from inactivation significantly faster in type B than type A neurons, and the recovery was dependent on conditioning voltage. These results suggest that while the rate of I(Na) inactivation is not responsible for the difference in the adaptation behavior of these two neuron types, the rate of recovery from inactivation may play a major role. Inactivation at lower potentials could therefore be crucial for more rapid recovery.

Predicting the responses of mechanoreceptor neurons to physiological inputs by nonlinear system identification.

The nonlinear dynamic properties of action potential encoding were studied in mechanosensory neurons innervating the slits of a slit-sense organ in the tropical wandering spider, Cupiennius salei. The organ contains two types of neurons that are morphologically similar but have different dynamic properties. Type A neurons produce only one or two action potentials in response to a mechanical or electrical stimulus of any suprathreshold amplitude, while type B neurons can fire prolonged bursts of action potentials in response to similar stimuli. Neurons were stimulated with pseudorandomly modulated intracellular current while recording the resultant fluctuations in membrane potential and action potentials. A parallel cascade method was used to estimate a third-order Volterra series to describe the nonlinear dynamic relationship between membrane potential and action potentials. Kernels measured for the two types of neurons had reproducible forms that showed differences between the two neuron types. The measured kernels were able to predict the responses of the neurons to novel pseudorandomly modulated inputs with reasonable fidelity. However, the Volterra series did not adequately predict the difference in responses to step depolarizations.

Low-voltage-activated calcium current does not regulate the firing behavior in paired mechanosensory neurons with different adaptation properties.

Low-voltage-activated Ca(2+) currents (LVA-I(Ca)) are believed to perform several roles in neurons such as lowering the threshold for action potentials, promoting burst firing and oscillatory behavior, and enhancing synaptic excitation. They also may allow rapid increases in intracellular Ca(2+) concentration. We discovered LVA-I(Ca) in both members of paired mechanoreceptor neurons in a spider, where one neuron adapts rapidly (Type A) and the other slowly (Type B) in response to a step stimulus. To learn if I(Ca) contributed to the difference in adaptation behavior, we studied the kinetics of I(Ca) from isolated somata under single-electrode voltage-clamp and tested its physiological function under current clamp. LVA-I(Ca) was large enough to fire single action potentials when all other voltage-activated currents were blocked, but we found no evidence that it regulated firing behavior. LVA-I(Ca) did not lower the action potential threshold or affect firing frequency. Previous experiments have failed to find Ca(2+)-activated K(+) current (I(K(Ca))) in the somata of these neurons, so it is also unlikely that LVA-I(Ca) interacts with I(K(Ca)) to produce oscillatory behavior. We conclude that LVA-Ca(2+) channels in the somata, and possible in the dendrites, of these neurons open in response to the depolarization caused by receptor current and by the voltage-activated Na(+) current (I(Na)) that produces action potential(s). However, the role of the increased intracellular Ca(2+) concentration in neuronal function remains enigmatic.

Temperature sensitivity of transduction and action potential conduction in a spider mechanoreceptor.

Previous work has suggested that the activation energy of mechanotransduction is higher than expected from the simple electrochemistry of ion channels, but the temperature sensitivity of mechanically activated receptor current has not been measured directly before. We used the single-electrode voltage-clamp technique to measure receptor currents in sensory neurons of the VS-3 slit-sense organ in the spider, Cupiennius salei. Receptor currents were generated by deforming the cuticular slits. Conduction velocity in afferent axons from the same organ was also measured by recording action potentials at two locations in the leg during mechanical stimulation of the slits. Activation energies of mechanotransduction and conduction velocity were estimated by making the measurements at a range of temperatures. The mean activation energy for receptor current was 23.1 kcal/mol (96.6 kJ/mol), corresponding to a Q10 value of 3.2. Conduction velocity in the afferent axons was approximately equal to 5 m/s at room temperature and it was much less temperature sensitive, with an activation energy of 6.3 kcal/mol (26.3 kJ/mol), corresponding to a Q10 value of 1.5. These results provide the first direct measurements of the activation energy of mechanically activated currents and support previous suggestions that a high thermal energy barrier is involved in mechanotransduction.

Hyposmotically activated chloride channels in cultured rabbit non-pigmented ciliary epithelial cells.

1. We used whole-cell patch-clamp recording techniques and noise analysis of whole-cell current to investigate the properties of hyposmotic shock (HOS)-activated Cl- channels in SV40-transformed rabbit non-pigmented ciliary epithelial (NPCE) cells. 2. Under conditions designed to isolate Cl- currents, exposure of cells to hyposmotic external solution reversibly increased the whole-cell conductance. 3. The whole-cell current activated with a slow time course (> 15 min), exhibited outward rectification and was Cl- selective. 4. The disulphonic stilbene derivatives 4, 4'-diisothiocyanatostilbene-2,2'-disulfonic acid (DIDS, 0.5 mM), 4-acetamido-4'-isothiocyanatostilbene-2,2'-disulfonic acid (SITS, 0. 5 mM) and 4,4'-dinitrostilbene-2,2'-disulfonic acid (DNDS, 0.5 mM) produced a voltage-sensitive block of HOS-activated Cl- current at depolarized potentials, whereas niflumic acid produced a voltage-independent block of the current. 5. Under Ca2+-free conditions, HOS stimulation still reversibly activated the Cl- current, but the amplitude of current was reduced and the time course of current activation was slower compared with control (P < 0. 05). 6. The non-specific kinase inhibitor H-7 (100 microM), upregulated HOS-activated Cl- current amplitude in all cells tested (P < 0.05). 7. Noise analysis of whole-cell Cl- current indicated that cell swelling activated a high density of small conductance Cl- channels (< 1 pS). 8. We conclude that HOS primarily activates a high density of volume-sensitive small conductance Cl- channels in rabbit NPCE cells, and that Ca2+ and phosphorylation are involved in channel regulation.

Primary culture of antennal mechanoreceptor neurons of Manduca sexta.

We have developed a primary cell culture system of antennal mechanoreceptor neurons from early-stage pupal sphinx moth Manduca sexta. Dissociated neurons from the moth antennae differentiated, grew and survived for several weeks in a conditioned culture medium. Bipolar neurons with soma diameters of 10-25 microns from the basal portion of the antennae could be positively identified as mechanoreceptor neurons, presumably derived from Johnston's organ, using a monoclonal antibody that recognizes neurofilaments in these neurons. The immunoreactivity was clear and specific from the first day after dissociation and became stronger during several days in culture. These neurons appeared healthy and showed normal whole-cell properties only a few days after plating. We found numerous mechanosensitive ion channels responding to both negative and positive pressures on the somata and neurites of differentiated neurons. This new culture system provides access to mechanoreceptor neurons that has never been possible before, allowing the use of both mechanical and electrical stimuli on neurons that are free from the accessory structures surrounding them in intact preparations.

Voltage-activated potassium outward currents in two types of spider mechanoreceptor neurons.

We studied the properties of voltage-activated outward currents in two types of spider cuticular mechanoreceptor neurons to learn if these currents contribute to the differences in their adaptation properties. Both types of neurons adapt rapidly to sustained stimuli, but type A neurons usually only fire one or two action potentials, whereas type B neurons can fire bursts lasting several hundred milliseconds. We found that both neurons had two outward current components, 1) a transient current that activated rapidly when stimulated from resting potential and inactivated with maintained stimuli and 2) a noninactivating outward current. The transient outward current could be blocked by 5 mM tetraethylammonium chloride, 5 mM 4-aminopyridine, or 100 microM quinidine, but these blockers also reduced the amplitude of the noninactivating outward current. Charybdotoxin or apamin did not have any effect on the outward currents, indicating that Ca2+-activated K+ currents were not present or not inhibited by these toxins. The only significant differences between type A and type B neurons were found in the half-maximal activation (V50) values of both currents. The transient current had a V50 value of 9. 6 mV in type A neurons and -13.1 mV in type B neurons, whereas the V50 values of noninactivating outward currents were -48.9 mV for type A neurons and -56.7 mV for type B neurons. We conclude that, although differences in the activation kinetics of the voltage-activated K+ currents could contribute to the difference in the adaptation behavior of type A and type B neurons, they are not major factors.

Principal dynamic mode analysis of nonlinear transduction in a spider mechanoreceptor.

The nonlinear dynamics of the mechanoelectrical transduction in an arthropod mechanoreceptor (cuticular slit sense organ of the spider Cupiennius salei) were studied using Volterra kernel measurements and the recently proposed method of principal dynamic modes. Since mechanoreceptors must operate with sufficient gain sensitivity to rapidly varying displacement stimuli over a broad bandwidth and for a wide range of amplitudes, the experimental data were generated by applying pseudorandom broadband mechanical displacements of various mean levels to the cuticular slits. The recorded response data were intracellular current and potential. The purpose of this paper is to demonstrate the use of the principal dynamic mode (PDM) methodology in elucidating the nonlinear dynamics of a spider mechanoreceptor. The results obtained demonstrate that two PDMs suffice to provide a complete nonlinear dynamic model of this insect mechanoreceptor. The first PDM resembles the first-order kernel and has a low pass characteristic (position dependent), while the second PDM has a high-pass characteristic (velocity-dependent) and resides entirely in the second-order kernel (nonlinear adaptation). This study may serve as an example of the proper use of this new methodology for the analysis of nonlinear physiological systems.

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.

Evaluation of the potential immunotoxicity of bromodichloromethane in rats and mice.

In the past two decades, concern has been expressed over the potential carcinogenicity of disinfection by-products (DBPs) found in chlorinated drinking water. More recently, research efforts have expanded to include noncancer endpoints as well. The objective of the present studies was to evaluate the potential of bromodichloromethane (BDCM), one of the most prevalent DBPs, to adversely affect immune function in mice and rats following drinking water or gavage exposure. Antigen-specific immunity was assessed as the antibody response to sheep erythrocytes; responses to T- and B-cell mitogens were evaluated as a non-antigen-specific measure of the proliferative potential of splenic and mesenteric lymph node lymphocytes. In consideration of an exposure route relevant to humans, C57BL/6 mice received 0.05, 0.25, or 0.5 g BDCM/L and F344 rats received 0.07 or 0.7 g BDCM/L via drinking water. In order to evaluate the effects of higher doses, animals were administered 50, 125, or 250 mg BDCM/kg/d (mice) or 75, 150, or 300 mg BDCM/kg/d (rats) via gavage. Under the conditions of these studies, no significant adverse effects on immune function were observed in mice. Despite some changes that were observed in non-antigen-specific immunity in rats, these experiments suggest that the immune system is not a sensitive target organ for BDCM toxicity.

Na+-Dependent neuritic spikes initiate Ca2+-dependent somatic plateau action potentials in insect dorsal paired median neurons.

The origin of plateau action potentials was studied in short-term cultures of dorsal paired median (DPM) neurons dissociated from the terminal abdominal ganglion of the cockroach, Periplaneta americana. Spontaneous plateau action potentials were recorded by intracellular microelectrodes in cell bodies that had neurite stumps. These action potentials featured a fast initial depolarization followed by a plateau. However, only fast spikes of short duration were observed when the cell was hyperpolarized from the resting membrane potential. These two different components of the action potentials could be separated by applying depolarizing current pulses from a hyperpolarized holding potential. Application of 200 nM tetrodotoxin (TTX) abolished both fast and slow phases, but depolarization to the original resting potential by steady current injection triggered slow monophasic action potentials that could be blocked by 3 mM CoCl2. In contrast, DPM neurons without neurites were not spontaneously active. In these cells, calcium-dependent slow monophasic action potentials were only recorded immediately after impalement or with current pulse stimulation. Immunocytochemical observations showed that dorsal unpaired median (DUM) neuron cell bodies, which are known to exhibit spontaneous sodium-dependent action potentials, reacted with an antibody directed against a synthetic peptide corresponding to the SP19 segment of voltage-activated sodium channels. In contrast, the antibody did not stain DPM neuron cell bodies but gave intense, patchy staining only in the neurite. Whole cell patch-clamp experiments performed on isolated DPM neuron cell bodies without a neurite revealed the presence of an inward current that did not inactivate completly within the duration of the test pulse. This current was insensitive to both 100 nM TTX and sodium-free saline. It was defined as a high-voltage-activated calcium current according to its high threshold of activation (-30 mV) and its sensitivity to 1 mM CdCl2 and 100 nM omega-conotoxin GVIA. Our findings demonstrate that spontaneous sodium-dependent spikes arising from the neurite are required to initiate slow somatic calcium-dependent action potentials in DPM neurons.

Adaptation properties of two types of sensory neurons in a spider mechanoreceptor organ.

The VS-3 slit-sense organs of the tropical wandering spider Cupiennius salei contain two types of mechanosensory neurons with similar morphology but different adaptation properties. We measured the changes in membrane potential produced by mechanical stimulation and by electric current stimulation in a large number of neurons of both types. No significant differences were found between the passive membrane properties of the two groups, but there were significant differences in the extent and time course of receptor potential adaptation between the two types of neurons. These data, combined with the responses to suprathreshold electrical stimuli, indicate that adaptational differences exist at several stages in these neurons but that active membrane conductances dominate the overall behavior. The passive membrane measurements also indicate that effective voltage clamp of the receptor current at the tips of the sensory dendrites is possible in these neurons.

Evaluation of the potential immunotoxicity of chlorinated drinking water in mice.

Recent epidemiological studies have reported associations between the consumption of chlorinated drinking water and various types of human cancer; in addition, exposure to chlorine (Cl-) in drinking water has been reported to suppress certain immune functions in laboratory animals. The current studies were conducted to extend our knowledge of the effects of drinking water exposure to Cl-. Female C57BL/6 mice were administered hyperchlorinated drinking water (7.5, 15, or 30 ppm Cl-) for 2 weeks prior to sacrifice for evaluation of spleen and thymus weights, the plaque-forming cell (PFC) response, hemagglutination (HA) titer, and lymphocyte proliferation (LP). Significant reductions in organ weights and immune response were observed in the positive control groups (i.e. dexamethasone- or cyclophosphamide-exposed mice). No consistent differences were observed between the Cl--exposed animals and vehicle control mice for the evaluated parameters. Thus, under the conditions of these experiments, 2 weeks of exposure to hyperchlorinated drinking water had no apparent adverse effects on immune function.

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.