RA Units Research Articles

Kolumbo submarine volcano (Greece): An active window into the Aegean subduction system.

Submarine volcanism represents ~80% of the volcanic activity on Earth and is an important source of mantle-derived gases. These gases are of basic importance for the comprehension of mantle characteristics in areas where subaerial volcanism is missing or strongly modified by the presence of crustal/atmospheric components. Though, the study of submarine volcanism remains a challenge due to their hazardousness and sea-depth. Here, we report 3He/4He measurements in CO2–dominated gases discharged at 500 m below sea level from the high-temperature (~220 °C) hydrothermal system of the Kolumbo submarine volcano (Greece), located 7 km northeast off Santorini Island in the central part of the Hellenic Volcanic Arc (HVA). We highlight that the mantle below Kolumbo and Santorini has a 3He/4He signature of at least 7.0 Ra (being Ra the 3He/4He ratio of atmospheric He equal to 1.39×10−6), 3 Ra units higher than actually known for gases-rocks from Santorini. This ratio is also the highest measured across the HVA and is indicative of the direct degassing of a Mid-Ocean-Ridge-Basalts (MORB)-like mantle through lithospheric faults. We finally highlight that the degassing of high-temperature fluids with a MORB-like 3He/4He ratio corroborates a vigorous outgassing of mantle-derived volatiles with potential hazard at the Kolumbo submarine volcano.

Development predictive QSAR models for artemisinin analogues by various feature selection methods: A comparative study

Quantitative structure–activity relationship (QSAR) models were derived for 179 analogues of artemisinin, a potent antimalarial agent. The activities of these compounds were investigated by means of multiple linear regression (MLR). To select relevant descriptors, several methods including stepwise selection, successive projection algorithm and an ant colony optimization algorithm (called memorized_ACS) were employed. A wide variety of molecular descriptors belonging to various structural properties were calculated for each molecule. Two matrixes (D1 and D2) of molecular properties were built. The D1 matrix included the calculated descriptors and the D2 matrix contained the first to third orders of the calculated descriptors and the logarithm of absolute values of the calculated descriptors. For both data matrixes, significant QSAR models were obtained by the memorized_ACS algorithm. The reactive and PEOE (partial equalization of orbital electronegativity) descriptors represented the highest impact on the antimalarial activity. The PEOE descriptors belong to partial charge descriptors and the reactive descriptor is an indicator of the presence of the reactive groups in the molecule. The best MLR model has a training error of 0.71 log RA units (r 2 = 0.81) and a prediction error of 0.48 log RA units (r 2 = 0.88).

Response properties of mouse trigeminal ganglion neurons

We used controlled whisker deflections to examine the response properties of 208 primary afferent neurons in the trigeminal ganglion of adult mice. Proportions of rapidly adapting (RA, 47%) and slowly adapting (SA, 53%) neurons were equivalent, and most cells had low or no spontaneous activity. We quantified angular tuning and sensitivity to deflection amplitude and velocity. Both RA and SA units fired more frequently to larger deflections and faster deflections, but RA units were more sensitive to differences in velocity whereas SA units were more sensitive to deflection amplitudes. Almost all neurons were tuned for deflection angle, and the average response to the maximally effective direction was more than fourfold greater than the average response in the opposite direction; SA units were more tuned than RA units. Responses of primary afferent whisker-responsive neurons are qualitatively similar to those of the rat. However, average firing rates of both RA and SA neurons in the mouse are less sensitive to differences in deflection velocity, and RA units, unlike those in the rat, display amplitude sensitivity. Subtle observed differences between mice and rats may reflect greater mechanical compliance in mice of the whisker hairs and of the tissue in which they are embedded.

Human tactile perception as a standard for artificial tactile sensing—a review

In this paper, we examine the most important features of human skin tactile properties with special emphasis on the characteristics which are vital in the design of artificial systems. Contrary to the visual and auditory senses, the touch signal is not a well-defined quantity. As a result, the researchers of this field are still dealing with the basics of collecting the most relevant data. Following this, mimicking the sense of touch by producing artificial tactile skin is a challenging process. Although the sense of touch is widely distributed all over the human body, the tactile perception in the human hand is of great importance in terms of surgical and medical robotics applications. In this study, the role of various mechanoreceptors in the human hand, such as, RA, SA I, SA II, and PC units are discussed in relation to the stimuli like force, position, softness, and surface texture. Taking human hand as a suitable tactile model, the necessary engineering features of an artificial tactile sensor, such as, spatial and temporal resolutions, force sensitivity, and linearity, are being reviewed. In this work, we also report on the current and possible future applications of tactile sensors in various surgical procedures.

Development of QSAR models to predict and interpret the biological activity of artemisinin analogues.

This work presents the development of Quantitative Structure-Activity Relationship (QSAR) models to predict the biological activity of 179 artemisinin analogues. The structures of the molecules are represented by chemical descriptors that encode topological, geometric, and electronic structure features. Both linear (multiple linear regression) and nonlinear (computational neural network) models are developed to link the structures to their reported biological activity. The best linear model was subjected to a PLS analysis to provide model interpretability. While the best linear model does not perform as well as the nonlinear model in terms of predictive ability, the application of PLS analysis allows for a sound physical interpretation of the structure-activity trend captured by the model. On the other hand, the best nonlinear model is superior in terms of pure predictive ability, having a training error of 0.47 log RA units (R2 = 0.96) and a prediction error of 0.76 log RA units (R2 = 0.88).

Novel information on peripheral tactile mechanisms in man acquired with concentric needle electrode microneurography

Microneurography with tungsten electrodes has provided a wealth of new data on peripheral nerve fibre function in man. Yet, some lingering controversies pertaining to the technique and its results have not been resolved. In particular, the working principles of microneurography allowing single unit sampling in man are not fully understood. Additionally debated, especially during recent years, was the validity of some neurographic data which supported the long standing conventional concept that myelinated fibres are randomly distributed intraneurally. A novel approach to address these issues was provided by microneurography with concentric needle electrodes. Data obtained with the latter technique suggested that these electrodes record activity extraaxonally from single myelinated fibres in man, possibly at or close to a node of Ranvier. The mechanisms described, which allow single unit resolution in humans, might well also be valid when performing microneurography with tungsten electrodes. Other sets of data indicated that Ranvier nodes tend to occur in clusters within certain regions of a nerve fascicle. Interestingly, the nerve fibres belonging to these clustering nodes were of the same modality and tended to innervate the same skin area in the hand. The discovered nerve fibre segregation involved all the four main classes of myelinated low threshold skin afferents in the hand (RA, PC, SAI and SAII units). The fact that sensory nerve fibres with clustering nodes and of the same modality tend to run together suggests at least a partially ordered intrafascicular nerve fibre organisation. The demonstrated intraneural fibre systematisation could be of profound functional significance both under normal conditions and in disease

Cutaneous sensory receptors in the rat foot

1. A total of 574 cutaneous afferent units in the sural and plantar nerves supplying the skin of the rat foot was examined: 399 A beta-units, 55 A delta-units, and 120 C-units. Their receptive-field (RF) properties were similar to those described in other mammals. However, the receptor type composition of units was different between the two nerves. 2. The sural A beta-fiber sample (n = 160) consisted of G-hair (41%), field (11%), rapidly adapting (RA; 6%), slowly adapting type I (SA-I; 7%), and type II (SA-II; 35%) mechanoreceptors. The plantar A beta-fiber sample (n = 239) was composed of G-hair (3%), RA (35%), SA-I (30%), SA-II (24%), and Pacinian corpuscle (PC; 8%) mechanoreceptors. 3. The RFs of SA-II units were located on both hairy and glabrous skin overlying the foot joints. Many of the SA-II units responded to movement of the foot joints. The RFs of both SA-I and RA units were small in size and located in high density on the toe tips and footpads. PC units were very sensitive to vibration and had extremely large RFs as in other species, although they were rare and found only in the plantar nerve. Field units were similar to SA-II units in response properties and RF distribution. 4. The sural A delta-fiber sample (n = 44) included nociceptors (68%), D-hair (27%), and cold (5%) receptors. All sampled plantar A delta-fibers (n = 11) were nociceptors. Of A delta-nociceptor units, A delta-mechanical nociceptors (73%) were dominant. 5. The sural C-fiber sample (n = 85) included nociceptors (44%), C-mechanoreceptors (33%), and cold receptors (21%). The plantar C-fiber sample (n = 35) included nociceptors (77%) and cold receptors (23%). No warm units were found among either the sural or plantar nerve fibers. Of C-nociceptors, C-mechanoheat nociceptors (80%) were dominant. 6. The results indicate that all well-known types of cutaneous receptors, except warm receptors, exist in the foot skin of the rat. On the basis of the fact that RFs of RA and SA-I units are in high density on the toe tips and footpads, it is suggested that those regions may have a spatial discriminating capacity. It is also suggested that SA-II receptors may play a role in proprioception, because they have RFs on the skin over foot joints and respond to joint movement.(ABSTRACT TRUNCATED AT 400 WORDS)

The raccoon lateral cervical nucleus: a single-unit analysis

1. Properties of 90 lateral cervical nucleus (LCN) neurons responsive to light tactile stimulation of ipsilateral body surfaces were examined in pentobarbital sodium-anesthetized raccoons. Peripheral receptive fields (RFs) of 60 of these lay totally or partially on glabrous skin of the forepaw. There were 71 neurons antidromically activated from the contralateral thalamic ventro-basal complex (VB) or medial lemniscus. Results were compared with previous findings in the raccoon spinocervical tract (SCT) and the dorsal column-medial lemniscal system (DC-MLS). 2. RFs located on glabrous skin of the digits were significantly smaller than those located on glabrous skin of the palm. All RFs, whether on glabrous skin of the forepaw or elsewhere, tended to be larger than those of either SCT or DC-MLS neurons. LCN units with glabrous forepaw RFs tended to be located ventrally within the nucleus. 3. Of those LCN neurons for which the RFs lay totally or partially on glabrous skin of the forepaw, relative numbers that were rapidly adapting (RA; 77%) versus slowly adapting (SA; 23%) were comparable with those found in the SCT and in VB. Relative numbers of LCN neurons that were classed as light touch (87%) versus multireceptive (13%) were comparable with those found in the SCT. 4. In contrast to both the SCT and VB, but in common with the prethalamic DC-MLS, indentation velocity coding functions of both RA and SA units fell within homogeneous groupings, power function exponents for RA units tending to be steeper than those for SA units (range of b = 0.710 - 0.919 vs. 0.448 - 0.883). 5. It is concluded that the raccoon spinocervicothalamic system (SCTS) as a whole lacks the "modality and place specificity" associated with the DC-MLS. Although the SCTS probably makes significant contributions to properties of VB neurons, these properties primarily reflect those of neurons of the DC-MLS.

Evidence for dual function in single RA units: [formula omitted] and H.E. Torebjörk (Dept. of Clin. Neurophysiol., Akademiska Sjukhuset, S-751 85 Uppsala, Sweden)

Evidence for dual function in single RA units: [formula omitted] and H.E. Torebjörk (Dept. of Clin. Neurophysiol., Akademiska Sjukhuset, S-751 85 Uppsala, Sweden)

Liminal and supraliminal response characteristics of mechanoreceptive neurons in the cuneate nucleus of cat.

The response characteristics of mechanoreceptive neurons (RA, SA, and PC) innervating the foot pad of cat were determined in the cuneate nucleus. The mechanical stimuli were single sinusoidal pulses of varying frequency (20, 60, 150, and 240 Hz), and vibratory trains of varying frequency (80 and 240 Hz) and duration (50, 100, and 400 ms). Thresholds and stimulus-response functions were determined with single pulses. Absolute thresholds (1 impulse/train), tuning thresholds (1 impulse/cycle), and atonal intervals (the range between absolute and tuning thresholds) were determined with vibratory stimulus trains. When tested with single pulses the thresholds resembled those of primary afferents in all unit populations. The stimulus-response function of PC units but not of all RA units were comparable to those of primary afferents. Noxious conditioning stimulation did not influence the thresholds of cuneate mechanoreceptors in the tested sample (N = 6). Mostly PC units were tested with vibratory trains. Absolute thresholds were not dependent on stimulus duration, which is a consistent finding with peripheral units. In contrast to peripheral units, the tuning thresholds in most PC units were elevated with increasing stimulus duration. The variability in the range of atonal intervals was much larger than in the periphery. Thus, it seems that both the type of the tactile signal and the type of the studied mechanoreceptive neuron are critical parameters in determining whether the response characteristics of neurons in the cuneate and in the periphery are identical or not.

Peripheral projections of nerve fibres in the human median nerve

Elementary sensations of definable quality were evoked by intraneural electrical stimulation of sensory units through tungsten microelectrodes inserted in the median nerve of awake human subjects. The most commonly reported sensations were tapping, pressure, dull pain and sharp pain, which correlated with recording from RA units, SA I units, C nociceptors and Aδ nociceptos, respectively. All sensations were projected more often to the fingertips than to the palm. The multifocal projections of sensations recruited by excitation of small groups of neighbouring nerve fibres were significantly more scattered in space than anticipated if these fibres had preserved an orderly intraneural map, implying lack of somatotopy in the nerve. Mean projected field sizes were 3.4 mm 2 for sharp pain, 9.2 mm 2 for tapping/vibration, 29.3 mm 2 for pressure and 35.4 mm 2 for dull pain, and in addition the latter two showed a rising size gradient from fingertip to palm. Consideration of projected field size as a variable in discriminative touch suggests that input from RA units and Aδ nociceptors may be of importance in stimulus localization, while SA I units may play an additional role in two point discrimination.

Sensations evoked by intraneural microstimulation of single mechanoreceptor units innervating the human hand.

Intraneural microstimulation (i.n.m.s.) was performed in awake human volunteers, using tungsten micro-electrodes inserted into median and ulnar nerve fascicles supplying the skin of the hand. The same electrodes were used alternatively to record impulse activity from single nerve fibres at the i.n.m.s. sites. Monitoring occasionally, with a proximal electrode, the impulse traffic evoked by i.n.m.s. distally in the same fascicle, established that the stimulation procedure could be made selective enough to activate single myelinated fibres in isolation, while also permitting multifibre recruitment. Monitoring propagated impulses also established that i.n.m.s. of a single myelinated fibre supplying a low-threshold mechanoreceptor in the hand might evoke an elementary sensation. Such sensations were fully endowed with cognitive attributes amenable to psychophysical estimation: quality, magnitude and localized projection. Psychophysical tests were made during i.n.m.s. at intraneural sites where single-unit activity was recorded from classified RA, PC, SA I or SA II mechanoreceptors. Changes in excitability of the nerve fibre of an identified unit, induced by further i.n.m.s., certified that the recorded unit had been stimulated during psychophysical tests. Comparing physical location of the receptive field of a recorded unit and localization of the projected field of the corresponding elementary sensation, revealed that either predicted the other accurately. This further assisted identification of the unit activated by i.n.m.s. The type of a recorded unit and the quality of the elementary sensation evoked by its activation were also reciprocally predictive. RA units evoked intermittent tapping, PC units vibration or tickle and SA I units evoked pressure. SA II units evoked no sensation when activated in isolation. Afferent impulse frequency determined magnitude of pressure in the SA I system, and frequency of vibration in the PC system. In the RA system, stimulation frequency determined frequency of tapping-flutter-vibration sensation at relatively low rates and subjective intensity of sensation at high rates. These findings endorse the concept that quality of sensation is coded in specific sensory systems. Further, they provide novel evidence that sensory quality, magnitude and localization can be exquisitely resolved at cognitive levels on the basis of input initiated in a single mechanoreceptor unit.

Primary afferent units from the hairy skin of the rat hind limb

The properties of cutaneous units in the saphenous nerve of the rat have been surveyed. We studied 137 units with myelinated (A-fibre) axons conducting at 4-44 m/s. Of the A-fibre units 66% gave a rapidly adapting discharge following hair movement and could be classified into categories similar to those described previously in cat and rabbit. Two categories predominated: (1) D-hair units with slowly conducting axons and relatively large receptive fields that responded to slow movement of hairs, including down hairs when these were present and (2) G-hair units with larger axons and relatively small fields that were only excited by fast movement of guard hairs. Of the A-fibre units 20% were high threshold mechanoreceptors. As in other species, these had a wide range of conduction velocities and multi-point receptive fields. Other types of A-fibre units found were (a) sensitive, RA units not excited by hair movement, (b) relatively insensitive RA units with diffuse receptive fields and (c) slowly adapting mechanoreceptor units. We studied 149 units with unmyelinated (C-fibre) axons conducting at 0.49-0.89 m/s. Of the C-fibre units 73% were of the polymodal nociceptor type. They had small receptive fields and responded to pressure and heating. The average heat threshold was 47 degrees C (+/- 6 degrees C, S.D.). Units were often not sensitized by suprathreshold heating unlike similar units in cat and rabbit. Other C-fibre units found were sensitive mechanoreceptors (12%), cold thermoreceptors (4%) or were very insensitive or inexcitable (11%). The pattern of innervation of rat limb hairy skin resembles previously studied mammalian species. A notable feature is the large proportion of C-polymodal nociceptor units. In this respect the rat resembles the primate and differs from the cat.

Sensations evoked from the glabrous skin of the human hand by electrical stimulation of unitary mechanosensitive afferents

A needle microelectrode in the median nerve was used to record impulses from tactile afferent units and to deliver trains of electrical pulses for microstimulation. The findings suggest that the activation of single SA I units produces a sensation of light pressure within a small area located within the receptive field of the unit whereas the activation of RA units seemed to elicit sensations of varying qualities: touch, vibration, and tickle.

Thresholds of mechanosensitive afferents in the human hand as measured with von Frey hairs

Afferent activity in low threshold mechanoreceptive units innervating the glabrous skin area of the hand was recorded from the median nerve of human subjects, using tungsten needle electrodes. The units were classified as RA, PC, SA I and SA II units mainly on the basis of their adaptation and receptive field properties Thresholds of the units were determined by two methods, the force threshold was determined with von Frey hairs and the indentation threshold with triangular indentations of controlled amplitudes. The characteristics of the force developed by the von Frey hairs as a function of movement parameters was analyzed when the hairs were prodded against a force transducer. The rise time was 10–20 msec and the variability of the peak force was within-20% of the steady-state force during repetitive prodding with various repetition rates. The von Frey thresholds were determined for 284 units and the indentation threshold for 116 of them. The RA and PC units were the most sensitive unit types with median thresholds of 0.58 mN and 0.54 mN. The medians of the slowly adapting SA I and SA II units, were 1.3 mN and 7.5 mN. The threshold distributions for all unit types were positively skewed. No regional differences were found but the thresholds were identical in skin areas where the psychophysical thresholds have been shown to be diverse. The relation between force threshold and indentation threshold was analyzed for 116 units. There was a positive correlation between the two thresholds, although the relation appeared not to be identical for the four types of units and the scatter was considerable.

Tactile sensibility in the human hand: receptive field characteristics of mechanoreceptive units in the glabrous skin area.

1. Impulses in tactile units innervating the glabrous skin of the hand recorded from the median nerve of adult human subjects. The recording electrodes which were made of tungsten were inserted percutaneously in the upper arm. 2. The units were classified on the basis of their sensitivity to sustained identation and to remote stimuli. Two types of rapidly adapting units and two types of slowly adapting units were found. In accordance with earlier reports they were denoted RA and PC units, and SA I and SA II units. 3. The sensitivity profiles of the receptive fields were analysed by measuring the extent of the receptive field as a function of the identation amplitude. 4. The RA and SA I units had receptive fields with several zones of maximal sensitivity distributed over an approximately circular or oval area typically covering five to ten papillary ridges. Within this area the sensitivity was high, whereas the sensitivity diminished steeply with increasing distance from this area. 5. The PC and SA II units had receptive fields with a single zone of maximal sensitivity and gentle continuous threshold increase outside this zone. 6. The relation between the identation amplitude and the receptive field size of a unit was described by a power function. The power exponent for the RA and SA I units was well below unity, whereas the PC and SA II units had exponenets greater than unity. The variation in exponent was very small among the RA and among the SA I units. Their average exponents were therefore regarded as characteristics of the unit type. 7. The SA II units were qualitatively studied with regard to their sensitivity to lateral skin stretch. Three different types of SA II units were described with regard to the directional pattern of this sensitivity. 8. The relation of the present findings to the problem of correlation between morphological structures of nerve end-organs and physiologically defined unit types is considered. 9. Moreover, the findings, indicate that the RA and SA I units are well suited for the analysis of mechanical events on the skin surface with a high degree of spatial selectivity, whereas the PC and SA II units are suited for analysis of other mechanical events, e.g. vibration and various forms of tension in the skin and related tissues.

Correlative physiological and morphological studies of rapidly adapting mechanoreceptors in cat's glabrous skin.

1. A total of fifty-four mechanoreceptor afferent units with fast conducting axons in the tibial nerve innervating the glabrous skin of the hind leg were isolated in anaesthetized cats. 2. Twenty-six rapidly adapting units (RA), eighteen slowly adapting units (SA) and ten Pacinian corpuscle units (PC) were differentiated from each other mainly on the presence of the off response in RA and PC units to a ramp stimulation, the persistence of discharges of the SA units during steady pressure on the receptive field and the classical tuning curve seen in the PC units. A few PC units in the hairy skin were also studied for comparison. 3. Lamellated corpuscles were found histologically in the skin of the receptive field of RA units and identified as Krause's corpuscle of cylindrical type by their superficial location in the cutaneous tissue and their structure revealed by electron microscopy. 4. Physiological characteristics of RA units to various forms of mechanical stimulation were studied and compared with those of the other two kinds of units. SA units had the lowest critical slope among three groups and PC units the highest. 5. The discharge pattern of RA and PC units to a ramp stimulation was found to be time-locked, whereas with SA unites only the first spike appeared at a fixed latency from the start of stimulation. 6. Some RA units showed a tuning curve which was flat from 10 to 200 Hz. Those with narrowly tuned curves had a best turning frequency at around 20 Hz. They were easily differentiated from the SA and PC units. SA units were tuned best at 5 HZ or less, and PC units at around 200 HZ. 7. The relation between the indentation velocity and amplitude of the ramp and the spike discharges was analysed in eleven RA units. In most cases the relation between identation velocity and maximum instataneous frequency was found to be best fit with a power function although other kinds of functions (linear, logarithmic, and logarithmic hyperbolic tangent) could also fit the relation at the 1% significance level. The instantaneous impuse frequency in RA units in response to various indentation amplitudes showed a step function. 8. The "off" responses to a ramp stimulation in RA units were also analysed in detail.

A deep 226Ra maximum in the Northeast Pacific

Seven vertical profiles of 226Ra have been measured along an east-west traverse at about 30°N from San Diego to northwest of Hawaii. These profiles show that there is a distinct core of Ra maximum spreading westward as a tongue in the northeast Pacific deep water. This core starts in the east with 21.1 Ra units (1Ra unit= 10 −14g/kg) at 3.9 km depth at about 130°W, and deepens westward to 4.1 km with its Ra reduced to 18.3 units at 150°W. A similar core with some uncertainty due to possible sampling errors extends westward near the bottom at 5.2 km depth from 19.4 Ra units at 150°W to 15.9 units at about 180° longitude. In addition, these profiles appear to be correlated with each other in structure above the cores of Ra maximum. These cores indicate that the Ra input depends locally on the type and composition of sediments and so the flux varies over the ocean bottom. On the basis of a one-dimensional diffusion-decay model, a horizontal diffusion coefficient of 10 6 cm 2/sec has been computed along these cores. Although this value appears to be slightly lower, it is not inconsistent with those derived from other physical methods.

Afferent impulses in median nerve fascicles evoked by tactile stimuli of the human hand

The median nerve mechanoreceptive innervation of the human hand was subjected to electrophysiological analysis with the microneurographic technique described by Vallbo and Hagbarth 45. Single-unit recordings from large axons impaled or damaged by the electrode tip revealed two main types of mechanoreceptive afferents with small, often ‘spot-like’ receptive fields in the glabrous skin: rapidly adapting touch units (RA units) and slowly adapting pressure units (SA units). Impulses in units of these two types were the main contributors to the composite multi-unit signals recorded extra-cellularly from individual median nerve fascicles in response to tactile stimuli within the fascicular innervation zones. The results indicate a large extracellular spread of unitary potentials within the fascicles with little or no ‘overhearing’ of the signals between adjacent fascicles. No even a maximal synchronous afferent volley in one fascicle could as a rule be ‘overheard’ by a microelectrode inserted in a neighbouring fascicle. Psychophysical and clinical applications of the microneurographic method are discussed.