The frequency selectivity of information-processing channels in the tactile sensory system

The frequency selectivity of the P, NP I, and NP II channels of the four-channel model of mechanoreception for glabrous skin was measured psychophysically by an adaptation tuning curve procedure. The results substantially extend the frequency range over which the frequency selectivity of these channels is known and further confirm the hypothesis that the input stage of each of these channels consists of specific sensory nerve fibers and associated receptors. Specifically, the frequency characteristics of Pacinian nerve fibers, rapidly adapting (RA) nerve fibers, and slowly adapting Type II (SA II) nerve fibers were found to be the peripheral neurophysiological correlates of the P, NP I, and NP II channels, respectively. The finding that the tuning characteristic for a test stimulus of 250 Hz delivered through a small (0.008 cm²) contactor depended dramatically on the duration of the test stimulus whereas the detection threshold did not, provides new evidence in support of the hypothesis that separate NP II and P channels exist.     

A four-channel analysis of the tactile sensitivity of the fingertip: frequency selectivity,spatial summation,and temporal summation

Thresholds were measured for the detection of vibratory stimuli of variable frequency and duration applied to the index fingertip and thenar eminence through contactors of different sizes. The effects of stimulus frequency could be accounted for by the frequency characteristics of the Pacinian (P), non-Pacinian (NP) I, and NP III channels previously determined for the thenar eminence (Bolanowski et al., J Acoust Soc Am 84: 1680-1694, 1988; Gescheider et al., Somatosens Mot Res 18: 191-201, 2001). The effect of changing stimulus duration was also essentially identical for both sites, demonstrating the same amount of temporal summation in the P channel. Although the effect of changing stimulus frequency and changing stimulus duration did not differ for the two sites, the effect of varying the size of the stimulus was significantly greater for the thenar eminence than for the fingertip. The attenuated amount of spatial summation on the fingertip was interpreted as an indication that the mechanism of spatial summation consists of the operations of both neural integration and probability summation.     

Spatial summation in the tactile sensory system: Probability summation and neural integration

Psychophysical thresholds for the detection of a 300-Hz burst of vibration applied to the thenar eminence were measured for stimuli applied to the skin through 1.5?cm² and through 0.05?cm² contactors. Thresholds were approximately 13?dB lower when the area of the contactor was 1.5?cm² than when it was 0.05?cm². The difference between the thresholds measured with the large and small contactors was significantly reduced when only the lowest thresholds obtained in the testing sessions were considered. This result supports the hypothesis that one component of spatial summation in the P channel is probability summation. In addition, threshold measurements within a session were less variable when measured with the 1.5?cm² contactor. We conclude that spatial summation in the P channel is a joint function of two processes that occur as the areal extent of the stimulus increases: probability summation in which the probability of exceeding the psychophysical detection threshold increases as the number of receptors of varying sensitivities increases, and neural integration in which neural activity originating from separate receptors is combined within the central nervous system rendering the channel more sensitive to the stimulus.     

Arthropod touch reception: structure and mechanics of the basal part of a spider tactile hair

Our previous studies on spider tactile hairs concentrated on the mechanical behavior of the hair shaft and the electrophysiological properties of the sensory cells. Here we focus on the structure and mechanical properties of the coupling of the hair shaft and the sensory terminals. 1. The functional design of the coupling provides for a combination of high sensitivity and protection against mechanical damage and overstimulation. The dendritic sheath is not directly coupled to the hair shaft. Rather, there is terminal connecting material between the dendrites and the hair shaft. 2. The hair shaft forms a first-order lever. Its acentric axis of rotation is located ca.3.5 m from its inner end. Displacement of the hair tip is scaled down by a factor of ca.750:1, not even considering the outer hair shafts bending. 3. At threshold the dendrite sheath displacement is ca. 0.05 m by forces in the order of 0.4–4×10^–6 N. 4. The hair shaft bends within the socket even before contacting it. The elasticities representing its suspension and bending in the socket can be described quantitatively by measuring the hairs restoring moments (range: 10^–9 Nm) and bending at different degrees of deflection.     

Perception of the tactile texture of raised-dot patterns: A multidimensional analysis

An ALSCAL multidimensional scaling analysis in Euclidean space revealed that three orthogonal perceptual dimensions can account for the judged tactile dissimilarities of raised-dot patterns. Through magnitude estimates of various perceptual attributes, it was determined that the three dimensions consist of blur, roughness, and clarity. The only effect that selective adaptation of the Pacinian (P) channel had was to change the perceptual clarity of the raised dots against their background. Adaptation of the P channel with a 20?dB SL 250?Hz stimulus enhanced clarity. As indicated by magnitude estimates, adaptation of the P channel by the 250?Hz stimulus had no effect on the perceived roughness of the dot pattern but did cause the individual dots of the textured pattern to feel smoother. When the observer was required to estimate magnitude “overall roughness” defined as a combination of dot-pattern roughness and individual-dot roughness, adaptation of the P channel affected perceived roughness by reducing it. Taken as a whole, the results are consistent with the hypothesis that the NP channels and the P channel jointly influence the perception of textured surfaces.     

Tactile sensitivity,tactile acuity,and affective touch: from childhood to early adolescence

The development of somatosensation and affective touch acquires a central role throughout our lives, for several reasons. In adults, these functions are driven by different, neuroanatomically and functionally segregated fibres. To date, very little is known about the basic features of these fibres in childhood and this lack of knowledge is mirrored in the affective touch domain, where there are no studies on the main physiological features of the tactile processes linked to the stimulation of the hairy skin, namely the preferential site of affective touch. Thus, our study aims to analyze (1) tactile sensitivity and tactile acuity of children’s hairy forearms; (2) a possible dissociation between somatosensation and the affective touch; and (3) the presence/absence of the perception of affective touch already in childhood. To these aims, participants (160 children, aged 6 to 14?years), were administered with the Von Frey (tactile sensitivity) and the 2 Point Discrimination (tactile acuity) tests. Affective touch was measured following the classic protocol and pleasantness ratings were recorded. Our findings showed a correlation between age and somatosensation, suggesting a progressive reduction of sensitivity and acuity as age grows. Further, there was no overlap between affective touch and somatosensation, suggesting a behavioural segregation. Lastly, we found higher pleasantness ratings for Affective versus Neutral stimulations at all ages and an enhanced preference for Affective as age grows. We concluded that both somatosensation and affective touch are already present as two separate components of touch in childhood and change as a function of age.     

Assessment of spatial acuity at the fingertip with grating (JVP) domes: validity for use in an elderly population

JVP domes are of a set of small grating surfaces recently introduced for cutaneous spatial resolution measurement. The gratings are placed on the skin and subjects are required to identify the orientation of grooves and bars. The finest grating whose orientations are discriminated reliably (75% correct) provides an estimate of the spatial resolution limit in the tested area. In the present study, we sought to determine the capacity of elderly subjects to resolve such grating stimuli in order to obtain normative data for this population. Thirty-two elderly individuals in good health (range: 60-88 years) were assessed for their ability to perceive grating orientation at the tip of the dominant index finger. Testing proceeded from the widest grating dome (3 mm) to the next (e.g., 2 mm), until the performance level dropped below 75% correct discrimination. The grating orientation task proved to be very difficult for most subjects and only a minority (14/32) was able to provide reliable reports of grating orientation even with presentation of the widest dome available (3 mm). Accordingly, individual grating resolution thresholds were often considerably higher (>2.5 mm, n = 26) than values previously reported in young adults for the fingertip region (approximately 1 mm). These results suggest that the current set of grating domes may not be adequate for spatial acuity measurement at the fingertip of older adults. New larger grating dimensions should be added to the set presently available to improve their sensitivity for an older population.     

Interaction of spatial and temporal summation in the warmth sense.

Two experiments explored the interaction of spatial and temporal summation in the perception of radiant heat. The first experiment showed that the absolute threshold for detecting two brief (0.1 sec) pulses of heat, one presented to each side of the forehead, is lower than the threshold for detecting a single pulse, even when the two pulses are separated in time by as muchas 0l75 sec. The second experiment showed that a single, 4-sec pulse of heat presented to one side of the forehead feels no warmer than two successive, 2-sec pulses, one to each side. Spatial and temporal summation appear to take place at least partly in the central nervous system.     

Possible roles of exceptionally conserved residues around the selectivity filters of sodium and calcium channels

In the absence of x-ray structures of sodium and calcium channels their homology models are used to rationalize experimental data and design new experiments. A challenge is to model the outer-pore region that folds differently from potassium channels. Here we report a new model of the outer-pore region of the NaV1.4 channel, which suggests roles of highly conserved residues around the selectivity filter. The model takes from our previous study (Tikhonov, D. B., and Zhorov, B. S. (2005) Biophys. J. 88, 184-197) the general disposition of the P-helices, selectivity filter residues, and the outer carboxylates, but proposes new intra- and inter-domain contacts that support structural stability of the outer pore. Glycine residues downstream from the selectivity filter are proposed to participate in knob-into-hole contacts with the P-helices and S6s. These contacts explain the adapted tetrodotoxin resistance of snakes that feed on toxic prey through valine substitution of isoleucine in the P-helix of repeat IV. Polar residues five positions upstream from the selectivity filter residues form H-bonds with the ascending-limb backbones. Exceptionally conserved tryptophans are engaged in inter-repeat H-bonds to form a ring whose π-electrons would facilitate passage of ions from the outer carboxylates to the selectivity filter. The outer-pore model of CaV1.2 derived from the NaV1.4 model is also stabilized by the ring of exceptionally conservative tryptophans and H-bonds between the P-helices and ascending limbs. In this model, the exceptionally conserved aspartate downstream from the selectivity-filter glutamate in repeat II facilitates passage of calcium ions to the selectivity-filter ring through the tryptophan ring. Available experimental data are discussed in view of the models.     

Meta-analysis,funnel plots and sensitivity analysis

Publication bias is a major problem, perhaps the major problem, in meta-analysis (or systematic reviews). Small studies are more likely to be published if their results are 'significant' than if their results are negative or inconclusive, and so the studies available for review are biased in favour of those with positive outcomes. Correcting for this bias is not possible without making untestable assumptions. In this paper, a sensitivity analysis is suggested which is based on fitting a model to the funnel plot. Some examples are discussed.     

A neuronal network model of primary visual cortex explains spatial frequency selectivity

We address how spatial frequency selectivity arises in Macaque primary visual cortex (V1) by simulating V1 with a large-scale network model consisting of O(10⁴) excitatory and inhibitory integrate-and-fire neurons with realistic synaptic conductances. The new model introduces variability of the widths of subregions in V1 neuron receptive fields. As a consequence different model V1 neurons prefer different spatial frequencies. The model cortex has distributions of spatial frequency selectivity and of preference that resemble experimental findings from the real V1. Two main sources of spatial frequency selectivity in the model are the spatial arrangement of feedforward excitation, and cortical nonlinear suppression, a result of cortical inhibition. Action Editor: Jonathan D. Victor     

A sensitivity analysis of the volumetric spatial decomposition algorithm

Recently, we proposed a new computational approach for the volumetric spatial decomposition of a three-dimensional bone structure into its basic rod and plate elements. This method was based on an image skeletonization approach, where two model parameters were used to identify an ideal skeleton. The goal of this study was to estimate the sensitivity of local morphometric indices to these two model parameters. Our results showed that the rod derived indices behaved more smoothly than plate derived indices, which suggests that rod derived indices are more trustworthy. The results also demonstrated that it was reasonable to reduce the model to only one parameter by setting the noise parameter n to twice the value of the slenderness parameter s. In conclusion, we found that local morphometric indices are reliable measures showing large differences between samples and thus may shed new light on structural differences of trabecular bone in a local fashion by adequately choosing one single optimization parameter.     

GENESIS: a simulation model of phylogeny 2. A sensitivity analysis

In a former paper (Heijerman 1988) a simulation model of phylogeny, GENESIS, was presented. This paer describes the results of a sensitivity analysis of GENESIS. The analysis is performed by cganging the input parameter values and estimating the relative effects on the model's output, as summarized by several tree statistics. The results show that none of the statistics tested can be classified as an unambiguous estimator of accuracy of methods for estimating hylogenetic trees. The sensitivity analysis increases the insight into the behaviour and applicability of the model. This is a prerequisite for a correct interpretation of the results of the evaluation experiments that will be carried out using GENESIS.     

Chemical derivatization of phosphoserine and phosphothreonine containing peptides to increase sensitivity for MALDI-based analysis and for selectivity of MS/MS analysis

Protein phosphorylation is one of the most important and common ways of regulating protein function in cells. However, phosphopeptides are difficult to analyse, ionising poorly under standard MALDI conditions. Several methods have been developed to deal with the low sensitivity and specificity of phosphopeptide analysis. Here, we show an approach using a simple one-step beta-elimination/Michael addition reaction for the derivatization of phosphoserine and phosphothreonine. The substitution of the negatively charged phosphate group by a positively charged S-ethylpyridyl group greatly improves the ionisation of the modified peptides, especially in MALDI MS, increasing the sensitivity of the analysis. The modification allows the formation of a unique fragment ion at m/z 106 under mild collisional activation conditions, which can be used for parent (precursor) ion scanning in order to improve both the sensitivity and the selectivity of the analysis. The optimisation of the approach is described for a standard model peptide and protein and then applied to phosphorylation analysis in two biologically derived proteins purified from different experimental systems.     

Modeling the mammalian circadian clock: sensitivity analysis and multiplicity of oscillatory mechanisms

We extend the study of a computational model recently proposed for the mammalian circadian clock (Proc. Natl Acad. Sci. USA 100 (2003) 7051). The model, based on the intertwined positive and negative regulatory loops involving the Per, Cry, Bmal1, and Clock genes, can give rise to sustained circadian oscillations in conditions of continuous darkness. These limit cycle oscillations correspond to circadian rhythms autonomously generated by suprachiasmatic nuclei and by some peripheral tissues. By using different sets of parameter values producing circadian oscillations, we compare the effect of the various parameters and show that both the occurrence and the period of the oscillations are generally most sensitive to parameters related to synthesis or degradation of Bmal1 mRNA and BMAL1 protein. The mechanism of circadian oscillations relies on the formation of an inactive complex between PER and CRY and the activators CLOCK and BMAL1 that enhance Per and Cry expression. Bifurcation diagrams and computer simulations nevertheless indicate the possible existence of a second source of oscillatory behavior. Thus, sustained oscillations might arise from the sole negative autoregulation of Bmal1 expression. This second oscillatory mechanism may not be functional in physiological conditions, and its period need not necessarily be circadian. When incorporating the light-induced expression of the Per gene, the model accounts for entrainment of the oscillations by light-dark (LD) cycles. Long-term suppression of circadian oscillations by a single light pulse can occur in the model when a stable steady state coexists with a stable limit cycle. The phase of the oscillations upon entrainment in LD critically depends on the parameters that govern the level of CRY protein. Small changes in the parameters governing CRY levels can shift the peak in Per mRNA from the L to the D phase, or can prevent entrainment. The results are discussed in relation to physiological disorders of the sleep-wake cycle linked to perturbations of the human circadian clock, such as the familial advanced sleep phase syndrome or the non-24h sleep-wake syndrome.     

A parameter sensitivity methodology in the context of HIV delay equation models

A sensitivity methodology for nonlinear delay systems arising in one class of cellular HIV infection models is presented. Theoretical foundations for a typical sensitivity investigation and illustrative computations are given.     

A probability-based analysis of temporal and spatial co-occurrence in grassland birds

Aim To test for non‐random co‐occurrence in 36 species of grassland birds using a new metric and the C‐score. The analysis used presence–absence data of birds distributed among 305 sites (or landscapes) over a period of 35 years. This analysis departs from traditional analyses of species co‐occurrence in its use of temporal data and of individual species’ probabilities of occurrence to derive analytically the expected co‐occurrence between paired species. Location Great Plains region, USA. Methods Presence–absence data for the bird species were obtained from the North American Breeding Bird Survey. The analysis was restricted to species pairs whose geographic ranges overlapped. Each of 541 species pairs was classified as a positive, negative, or non‐significant association depending on the mean difference between the observed and expected frequencies of co‐occurrence over the 35‐year time‐span. Results Of the 541 species pairs that were examined, 202 to 293 (37–54%) were positively associated, depending on which of two null models was used. However, only a few species pairs (<5%) were negatively associated. An additional 89 species pairs did not have overlapping ranges and hence represented de facto negative associations. The results from analyses based on C‐scores generally agreed with the analyses based on the difference between observed and expected co‐occurrence, although the latter analyses were slightly more powerful. Main conclusions Grassland birds within the Great Plains region are primarily distributed among landscapes either independently or in conjunction with one another. Only a few species pairs exhibited repulsed or segregated distributions. This indicates that the shared preference for grassland habitat may be more important in producing coexistence than are negative species interactions in preventing it. The large number of non‐significant associations may represent random associations and thereby indicate that the presence/absence of other grassland bird species may have little effect on whether a given focal species is also found within the landscape. In a broader context, the probability‐based approach used in this study may be useful in future studies of species co‐occurrence.     

A sensitivity analysis to methodological variation in indicator-based urban sustainability assessment: a Quebec case study

Methodological factors are often acknowledged for their impact on the urban sustainability assessment outcomes. The significance of this impact, however, divides those who assign it great importance; to the point of compromising any attempt at sustainability benchmarking using indicators, from those who regard it as having a relatively limited role. In this paper, we aim to contribute to this debate by assessing how methodological choices could influence the variation in the outcomes of indicator-based urban sustainability benchmarking. The paper starts with a literature review outlining the evolution of the approaches adopted since the pioneering experiences in the ‘90s to show a convergence in indicators and data over time. Subsequently, by using the data of the 25 most populous cities in Quebec, we empirically analyse the sensitivity of the cities’ sustainability scores to 18 combinations of estimation techniques, and observe consistency within the outcomes while maintaining general patterns regardless of the approach used. The results suggest that methodological choices play a limited role in the urban sustainability assessment outcomes. Thus, other determinants beyond the evaluation methods are most likely involved and therefore should be a major focus in future research agendas.     

Modelling C3 photosynthesis: A sensitivity analysis of the photosynthetic carbon-reduction cycle

A model of the C ₃ photosynthetic system is developed which describes the sensitivity of the steadystate rate of carbon dioxide assimilation to changes in the activity of several enzymes of the system. The model requires measurements of the steady-state rate of carbon dioxide assimilation, the concentrations of several intermediates in the photosynthetic system, and the concentration of the active site of ribulose 1,5-bisphosphate carboxyalse/oxygenase (Rubisco). It is shown that in sunflowers (Helianthus annuus L.) at photon flux densities that are largely saturating for the rate of photosynthesis, the steady-stete rate of carbon dioxide assimilation is most sensitive to Rubisco activity and, to a lesser degree, to the activities of the stromal fructose, 6-bisphosphatase and the enzymes catalysing sucrose synthesis. The activities of sedoheptulose 1,7-bisphosphatase, ribulose 5-phosphate kinase, ATP synthase and the ADP-glucose pyrophosphorylase are calculated to have a negligible effect on the flux under the high-light conditions. The utility of this analysis in developing simpler models of photosynthesis is also discussed.Abbreviations c _i intercellular CO₂ concentration - C _infP ^supJ control coefficient for enzyme P with respect to flux J - DHAP dihydroxyacetonephosphate - E4P erythrose 4-phosphate - F6P fructose 6-phosphate - FBP fructose 1,6-bisphosphate - FBPase fructose 1,6-bisphosphatase - G3P glyceraldehyde 3-phosphate - G1P glucose 1-phosphate - G6P glucose 6-phosphate - Pi inorganic phosphate - PCR photosynthetic carbon reduction - PGA 3-phosphoglyceric acid - PPFD photosynthetically active photon flux density - R _n ^J response coefficient for effector n with respect to flux J - R5P ribose 5-phosphate - Rubisco ribulose 1,5-bisphosphate carboxylase/oxygenase - Ru5P ribulose 5-phosphate - RuBP ribulose 1,5-bisphosphate - S7P sedoheptulose 7-phosphate - SBP sedoheptulose 1,7-bisphosphate - SBPase sedoheptulose 1,7-bisphosphatase - SPS sucrose-phosphate synthase - Xu5P xylulose 5-phosphate - _n ^P elasticity coefficient for effector n with respect to the catalytic velocity of enzyme P This research was funded by an Australian Research Council grant to I.E.W. and was undertaken during a visity by K.A.M. to the James Cook University of North Queensland. The expert help of Glenys Hanley and Mick Kelly is greatly appreciated.