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Spatial spread of activation and background desensitization in toad rod outer segments.

1. The spread of activation and background desensitization in rods was studied by recording membrane current from single outer segments in pieces of isolated toad retina.2. Flash sensitivity changed slightly along the outer segment, falling by about 30% from base to tip.3. When only the distal half of an outer segment was in the recording pipette, illumination of the unrecorded part elicited little or no photocurrent at the recorded part, indicating that a photoisomerization does not cause activation of the entire outer segment.4. With diffuse illumination of an outer segment fully drawn into the pipette, the intensity-response relations at fixed times were invariant in form for most of the rising phase of the flash response and were considerably steeper than the Michaelis relation. The observed relation was consistent with a model in which a photoisomerization blocks all channels over a short region of the outer segment.5. With illumination restricted to a narrow transverse slit, the intensity-response relations at fixed times were much less steep, as would be expected for a very limited longitudinal spread of activation. An upper limit for the effective longitudinal diffusion coefficient of the internal transmitter was estimated to be about 3 x 10(-7) cm(2) sec(-1). This corresponds to a space constant for longitudinal spread of transmitter of about 3 mum at the time of the dim response peak.6. The time course of flash responses elicited with light positioned either on the edge or on the centre of the outer segment was very similar.7. Desensitization resulting from steady illumination by a transverse slit was also localized longitudinally. A linear desensitization parameter T, defined in Results, decayed approximately exponentially along the outer segment, on either side of the site of photoisomerization, with a space constant of about 6 mum.8. Transverse spread of desensitization was more effective than longitudinal spread.9. After turning off a dim diffuse background light, the decay of T was roughly exponential with a time constant of several seconds. From this, and the steady state space constant of 6 mum, it is estimated that the effective longitudinal diffusion coefficient for a ;desensitizing substance' would also be about 10(-7) cm(2) sec(-1).10. The restricted longitudinal spread of activation and desensitization may be explained by the barrier to diffusion presented by the stacked membranous disks in the outer segment. This baffling reduces the effective longitudinal diffusion coefficient to about 1/50 that of ordinary aqueous diffusion, but it does not significantly affect transverse spread.

Animals

The effect of attentional spread on spatial resolution.

The effects of attentional spread were studied by having subjects detect a luminance increment along a row of evenly spaced dots. The increment could occur for the central, fixated dot (Narrow Attention) or for either the fixation dot or one of the four dots to its left or right (Broad Attention). Narrow Attention enhanced the detection of luminance increments for the fixated dot, and also enhanced spatial resolution near the fixation dot for judgments of vernier alignment and separation. This indicated that the sensitivity of small spatial filters in the fovea was increased more by narrowly focused than broadly spread attention. Effects of attentional spread on spatial resolution were not obtained for judgments of the separation between two peripherally located targets, perhaps because of their dependence on eccentricity (position) rather than separation.

Attention

The mass angular scattering power method for determining the kinetic energies of clinical electron beams.

A method for determining the kinetic energy of clinical electron beams is described. The method is based on the measurement in air of the spatial spread of a pencil electron beam which is produced from the broad clinical electron beam. As predicted by the Fermi-Eyges theory, the dose distribution measured in air on a plane, perpendicular to the incident direction of the initial pencil electron beam, is Gaussian. The square of its spatial spread is related to the mass angular scattering power which in turn is related to the kinetic energy of the electron beam. The measured spatial spread may thus be used to determine the mass angular scattering power, which is then used to determine the kinetic energy of the electron beam from the known relationship between mass angular scattering power and kinetic energy. Energies obtained with the mass angular scattering power method agree with those obtained with the electron range method. The angular scattering power method is relatively cumbersome, but allows us to determine the kinetic energies of electron beams from first principles, in contrast to the empirical methods based on range measurements in water.

Electrons

An in vivo pharmacological study of single group Ia fibre contacts with motoneurones in the cat spinal cord.

1. Direct experimental evidence was obtained on the spatial distribution of active synaptic contacts from single Ia muscle afferents on the dendrites of lumbosacral motoneurones in anaesthetized cats. 2. An extracellular micropipette was used to pressure eject the AMPA/kainate receptor antagonists 6-cyano-7-nitroquinoxaline-2,3-dione (CNQX) or 2,3-dihydroxy-6-nitro-7-sulphamoyl-benzo(F)quinoxaline (NBQX) in close proximity to the intracellular recording site, in order to create an extracellular concentration gradient of the antagonist. The effect of antagonist ejection on the time course and amplitude of excitatory postsynaptic potentials (EPSPs) evoked in motoneurones by impulses in single group Ia fibres was examined. 3. Pressure ejection of NBQX resulted in a complete block of the monosynaptic group Ia EPSP in two cells, and a significant reduction to 23-57% of control EPSP peak amplitudes in a further six cells (mean, 27%; n = 8). These effects were not associated with changes in membrane potential or membrane time constant. 4. The reduction in amplitude of these single group Ia fibre EPSPs following ejection of NBQX was usually accompanied by a pronounced slowing in the time course of the EPSPs. On average, the EPSP rise times and half-widths were increased by 269 and 37%, respectively. This is most probably due to a considerable spatial spread of the synaptic contacts along the dendrites of motoneurones, with the most proximal synaptic contacts (producing the briefest synaptic potentials) subjected to a greater reduction in amplitude due to a higher local antagonist concentration. 5. An equivalent dendritic cable model of the motoneurone was used to interpret the observed changes in the time course of single fibre EPSPs. The time course of control single fibre EPSPs examined in the present study could be well matched using the cable model and assuming a single location for synaptic input. The observation of a slowed EPSP time course following antagonist ejection indicated that this assumption was not correct and that there was in fact considerable spatial spread in the synaptic contacts arising from these single afferent fibres. These results provide direct evidence that spatial spread of synaptic input may not be detected using the time course of a synaptic potential in conjunction with a neuronal cable model of the postsynaptic cell.

6-Cyano-7-nitroquinoxaline-2,3-dione

The effect of angular spread on the intensity distribution of arbitrarily shaped electron beams.

Knowledge of the relative intensity distribution at the patient's surface is essential for pencil beam calculations of three-dimensional dose distributions for arbitrarily shaped electron beams. To calculate the relative intensity distribution, the spatial spread resulting from angular spread is convolved with a two-dimensional step function whose shape corresponds to the applicator aperture. Two different approaches to obtain angular spread or the equivalent spatial spread are investigated. In the first method, the pencil beam angular spread is assumed to be Gaussian in shape. The angular spread constants (sigma theta) are then obtained from the slopes of measured intensity profiles. In the second method, the angular spread, in the form of an array of numerical values, is obtained by the deconvolution of measured intensity profiles. After obtaining the angular spread, the calculation for convolution is done in a number of parallel planes normal to the central axis at various distances from the electron collimator. Intensity at any arbitrary point in space is computed by interpolating between intensity distributions in adjacent planes on either side of the point. The effects of variations in angular spread as a function of field size for two treatment machines, one with a scanned electron beam and the other with a scattering foil, have been studied. The consequences of assuming angular spread to be of Gaussian shape are also examined. The electron intensity calculation techniques described in this paper apply primarily to methods of dose calculations that employ pencil beams generated using Monte Carlo simulations.

Electrons

Spatiotemporal visual response to suprathreshold stimuli.

The spatiotemporal visual response to an inducing bar of a short duration was studied by a brightness-matching procedure. The apparent brightness of a test line, presented in the centre of an inducing bar, exhibited a U-shaped dependence of the inducing-bar width with a minimum at about 4.5' width. The temporal response to a 4.5'-wide inducing stimulus consisted of three alternating phases, the middle one being the largest. The spatial spread of the response to this stimulus was initially restricted in the area of the inducing bar, later it was extended to adjacent positions achieving a triphasic form and still later faded away. These findings indicate that the response to a spatiotemporal impulse of suprathreshold luminance is triphasic in time and the spatial spread of the response depends on the time after the stimulus onset. A model was presented assuming that the spatiotemporal weighting function might be described by a spherical harmonic function modulated by a Gaussian function. The model predictions agreed with the data obtained.

Contrast Sensitivity

Evidence that two sizes of ventromedial hypothalamic neurones project to the mesencephalic central grey matter in rats.

The spatial spread of the extracellular antidromic action potentials was measured in eighty-three neurones in the ventromedial nucleus region of the female rat hypothalamus following electrical stimulation of the mesencephalic central grey matter. The positive-negative configuration of the antidromic action potentials across the extracellular field suggested that the potentials were generated predominantly by neuronal soma with simple geometries. When represented by the distance travelled by the electrode, at which peak-to-peak spike amplitude exceeded the half maximum value, the spatial spread of the extracellular field ranged from 25 to 174 micron. The frequency distribution for the field size was distinctively bimodal and could be divided into large and small groups at 85 micron. Antidromic action potentials with larger extracellular fields had significantly larger maximum spike amplitude and shorter duration, indicating that differences in field size were associated with neuronal size. At least 55% of the central grey projection of the ventromedial nucleus originated from small neurones. Taking into account the sampling bias, a much greater proportion of the central grey projection may arise from small neurones. The lack of a systematic difference in the antidromic spike latencies between large and small cells indicated that axonal thickness is not the major factor in determining the latency of the responses of the ventromedial hypothalamic neurones to stimulation of the central grey matter.

Amygdala

Spatiotemporal impulse response and cortical magnification.

According to a model of the spatiotemporal weighting function (Manahilov, V. Spatiotemporal visual response at suprathreshold stimuli. Vision Research, 1995, 35, 227-237; and Triphasic temporal impulse responses and Mach bands in time. Vision Research, 38, 447-458) the waveform of the temporal-impulse response and the cortical spread of the spatial-impulse response should not depend on the retinal site of stimulation. To verify these model predictions, the spatiotemporal responses to brief near-threshold lines presented in the fovea and the near retinal periphery were studied. The effect of an inducing stimulus on the threshold for pattern detection of a test stimulus was measured, assuming that the pattern-detection threshold was determined by the test peak response. The spatial spread of the line response expressed in visual-field units was increased with eccentricity. The temporal-impulse responses to foveal and peripheral stimuli were similar. The model of the weighting function was used to evaluate the relative magnification factor for the retinal location tested. The calculated cortical spatial-impulse responses did not depend on the stimulation site. The data obtained are in line with the cortical magnification theory of peripheral vision.

Humans

Calcium inactivation of calcium release in frog cut muscle fibers that contain millimolar EGTA or Fura-2.

Cut muscle fibers from Rana temporaria (sarcomere length, 3.4-4.2 microns) were mounted in a double Vaseline-gap chamber (14-15 degrees C) and equilibrated with end-pool solutions that contained 20 mM EGTA and 1.76 mM Ca. Sarcoplasmic reticulum (SR) Ca release was estimated from changes in pH (Pape, P. C., D.-S. Jong, and W.K. Chandler. 1995. Journal of General Physiology. 106:000-000). Although the amplitude and duration of the [Ca] transient, as well as its spatial spread from the release sites, are reduced by EGTA, SR Ca release elicited by either depolarizing voltage-clamp pulses or action potentials behaved in a manner consistent with Ca inactivation of Ca release. After a step depolarization to -20 or 10 mV, the rate of SR Ca release, corrected for SR Ca depletion, reached a peak value within 5-15 ms and then rapidly decreased to a quasi-steady level that was about half the peak value; the time constant of the last half of the decrease was usually 2-4 ms. Immediately after an action potential or a 10-15 ms prepulse to -20 mV, the peak rate of SR Ca release elicited by a second stimulation, as well as the fractional amount of release, were substantially decreased. The rising phase of the rate of release was also reduced, suggesting that at least 0.9 of the ability of the SR to release Ca had been inactivated by the first stimulation. There was little change in intramembranous charge movement, suggesting that the changes in SR Ca release were not caused by changes in its voltage activation. These effects of a first stimulation on the rate of SR Ca release elicited by a second stimulation recovered during repolarization to -90 mV; the time constant of recovery was approximately 25 ms in the action-potential experiments and approximately 50 ms in the voltage-clamp experiments. Fura-2, which is able to bind Ca more rapidly than EGTA and hence reduce the amplitude of the [Ca] transient and its spatial spread from release sites by a greater amount, did not prevent Ca inactivation of Ca release, even at concentrations as large as 6-8 mM. These effects of Ca inactivation of Ca release can be simulated by the three-state, two-step model proposed by Schneider, M. F., and B. J. Simon (1988, Journal of Physiology. 405:727-745), in which SR Ca channels function as a single uniform population of channels. (ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials

The rod-cone shift and its effect on ganglion cells in the cat's retina.

We examined how several characteristics of cat retinal ganglion cells--receptive field size, spatial resolution, and centre-surround antagonism--change with background illumination. Spectral sensitivity was also measured to see how these changes depend on the rod-cone shift. The radius of the centre mechanism changed very little across the mesopic range. The absence of a change can be attributed to the connections rods make with cones, and to the small spatial spread of rods which connect to a cone. The highest spatial frequency to which a cell could respond dropped sharply with falling background illumination. This loss of spatial resolution is due partly to increasing receptive field size, and partly to loss of contrast gain. Centre-surround antagonism approached zero as background illumination fell. The loss of antagonism could have been due to either a change in the subtractive relationship between centre and surround, or due to a loss of surround strength relative to centre strength; the latter was shown to be the case.

Adaptation, Ocular

Topographic mapping of the visual evoked potential after source derivation.

Visual Evoked Potentials (VEP) have been considered an important complementary diagnostic method for the study of the physiology of the visual pathways and cortex. It is suggested that topographic maps of Visual Evoked Potentials (VEP) generate information related to the anatomy and physiology of the visual cortex. The source derivation technique applied to electroencephalographic signals has been shown to improve the sensitivity and spatial selectivity of the results. Mapping VEP after source derivation was implemented in order to facilitate the interpretation of the VEP signals and as an aid in identifying the distribution of the sources generating them. The signals were obtained from normal individuals stimulated with full-field and half-field pattern reversal checker-boards. The electrode system used (16 channels) was that proposed by Bodis-Wollner; this system concentrates the electrodes in the scalp areas related to the visual cortex. Signals were sampled simultaneously with a 10-bit A/D converter, saved and, thereafter, processed in an IBM/PC compatible microcomputer. After coherently averaging the responses, source derivation considering inter-electrode distances was applied. It was observed that the spatial spreading of the signals was reduced, leading to an improved spatial selectivity, locating the VEP sources in the expected brain regions and suggesting the existence of electrical dipoles.

Adult

The absence of spread of adaptation between rod photoreceptors in turtle retina.

Adaptation by weak backgrounds and the spatial spread of desensitization between rods was studied in the snapping turtle retina, Chelydra serpentina. Intracellular membrane potentials were recorded from these photoreceptors in an eyecup preparation. The kinetics and sensitivity of rod responses were changed significantly by large, very dim backgrounds. For the twenty-five most sensitive rods where the dark-adapted flash sensitivity, SDF, was greater than 1.0 mV/Rh*, Rh* being the number of effective photo-isomerizations per rod, the background intensity required to halve the amplitude of the linear range response averaged 0.21 Rh* s-1. The time-to-peak of the test responses was reduced up to 50% by these dim backgrounds. The desensitizing effects of full field backgrounds of various intensities on the responses to large test spots were measured. The dependence of incremental flash sensitivity, SF, on background intensity, IB, followed the form (FORMULA: SEE TEXT) where I0 is the background intensity which halved SDF. The same intensity dependence held for slit-shaped background fields that desensitized responses to small test spots. The desensitizing effects of large, very dim flashed and continuous backgrounds took several seconds to appear and decay to dark levels. This in conjunction with the sparsity of photons suggests, that the desensitization from a single photoisomerization can persist for several seconds. A comparison of the desensitizing effects of spot and annular backgrounds revealed that small spot backgrounds superimposed on the centered test spots desensitized rods more effectively than annular fields. This finding held true even when annular patterns produced a greater maintained hyperpolarization in the rods. Thus, there was no unique relationship between desensitization and the steady maintained hyperpolarization evoked by a background field. The dependence of adaptation on distance from the impaled rod was determined with slit-shaped background fields placed at different positions across the rod's receptive field. The desensitizing effect of displaced slit stimuli was found to decline much more rapidly with distance than excitation. Displacing the slit by 20 micron from the centre reduced its desensitizing effect by more than 1 log unit. In contrast, excitation fell to about 80% at the same distance (lambda ranging from 50 to 70 micron). The fall off of desensitization with distance matched the calculated fall off with distance of light scatter from a slit. No difference was noted in the kinetics of test responses in the presence of equally desensitizing, superimposed and displaced slits.(ABSTRACT TRUNCATED AT 400 WORDS)

Adaptation, Ocular

Electrical interactions via the extracellular potential near cell bodies.

Ephaptic interactions between a neuron and axons or dendrites passing by its cell body can be, in principle, more significant than ephaptic interactions among axons in a fiber tract. Extracellular action potentials outside axons are small in amplitude and spatially spread out, while they are larger in amplitude and much more spatially confined near cell bodies. We estimated the extracellular potentials associated with an action potential in a cortical pyramidal cell using standard one-dimensional cable theory and volume conductor theory. Their spatial and temporal pattern reveal much about the location and timing of currents in the cell, especially in combination with a known morphology, and simple experiments could resolve questions about spike initiation. From the extracellular potential we compute the ephaptically induced polarization in a nearby passive cable. The magnitude of this induced voltage can be several mV, does not spread electrotonically, and depends only weakly on the passive properties of the cable. We discuss their possible functional relevance.

Action Potentials

Comparison of methods to determine electron pencil beam spread in tissue-equivalent media.

This study has intercompared the predictions of Fermi-Eyges theory for the rms spatial spread (sigma) of an electron pencil beam scattering in muscle-, lung- and bone-equivalent media with those of; two range straggling modifications to the theory, Monte Carlo simulations, and an empirical method based on broad beam penumbra. Systematic differences among the results obtained by these methods for the values of sigma have been identified. Monte Carlo simulations are lower than the predictions of Fermi-Eyges theory for sigma at all depths whereas the broad beam penumbra method results are in reasonable agreement with Fermi-Eyges theory at depths less than approximately 0.7 times the range of the incident electrons. All of the methods investigated have an increasing discrepancy from the predictions of Fermi-Eyges theory with depth, especially close to the end of the electron range. The two range-straggling modifications to Fermi-Eyges theory developed for soft tissue do not agree with either measured or Monte Carlo results for sigma in homogeneous scattering media of lung and bone.

Bone and Bones

Nonlinear processing of tactile information in the thalamocortical loop.

Rats explore tangible objects in a manner such that, at any given moment in time, multiple facial whiskers simultaneously contact the surface of the object. Although both thalamic and cortical neurons responsible for processing such tactile information have large, multiwhisker receptive fields, it remains unclear what kinds of computations can be carried out by these neuronal populations when behaviorally relevant multiwhisker stimuli are used. By simultaneously recording the activity of up to 78 cortical and thalamic neurons per animal, we observed that the magnitude of sensory responses and the spatial spread of ensemble activity increased in a nonlinear fashion according to the extent and spatial orientation of the multiwhisker stimuli. Supralinear responses were seen more frequently with vertically than with horizontally oriented stimuli. These data suggest that thalamocortical interactions in the rat somatosensory system can generate complex spatial transformations of multiwhisker stimuli that go beyond the classic inhibitory interactions previously observed.

Animals

Estimating exposure to volatile organic compounds from municipal water-supply systems: use of a better computational model.

The Southington, Connecticut, water-supply system is characterized by a distribution network that contains more than 1 700 pipeline segments of varying diameters and construction materials, more than 186 mi (299 km) of pipe, 9 groundwater extraction wells capable of pumping more than 4 700 gal/min (0.2965 m3/s), and 3 municipal reservoirs. Volatile organic compounds, which contaminated the underlying groundwater reservoir during the 1970s, contaminated the water-supply system and exposed the town's residents to volatile organic chemicals. We applied a computational model to the water-supply system to characterize and quantify the distribution of volatile organic compounds in the pipelines, from which we estimated the demographic distribution of potential exposure to the town's residents. Based on results from modeling analyses, we concluded the following: (a) exposure to volatile organic compound contamination may vary significantly from one census block to another, even when these census blocks are adjacent to each other within a specified radius; (b) maximum spatial spread of contamination in a water-distribution system may not occur under peak demand conditions, and, therefore, maximum spatial distribution of the exposed population also may not correspond to peak demand conditions, and (c) use of the proposed computational model allows for a more refined and rigorous methodology with which to estimate census-block-level contamination for exposure assessment and epidemiologic investigations.

Algorithms

A spatial-frequency dependent quantum accounting diagram and detective quantum efficiency model of signal and noise propagation in cascaded imaging systems.

The detective quantum efficiency (DQE) is a system parameter that can be used to accurately describe image noise transfer characteristics through many imaging systems. A simpler approach used by some investigators, particularly when evaluating new ideas and system designs, is to describe the system as a series of cascaded stages. Each stage may correspond to either an increase in the number of quanta (e.g., conversion from x-ray to optical quanta in a radiographic screen), or a loss (a detection or coupling probability). The number of secondary quanta at each stage per incident primary quantum is given by the product of all preceding gains, and can be displayed graphically for convenient interpretation. The stage with the fewest quanta is called the "quantum sink," limiting the pixel signal-to-noise ratio to less than the square root of the number of quanta per pixel. This conventional zero-spatial-frequency "quantum accounting diagram" (QAD), however, neglects the spatial spreading of secondary quanta and can seriously underestimate image noise. It is shown that this problem is avoided with the introduction of a spatial-frequency dependent QAD, expressed as the product of the gains and squared modulation-transfer functions (MTF) of each stage. A generalized expression is developed for the DQE of a cascaded imaging system that is dependent only on the gain, gain Poisson excess (related to the variance), and MTF, of each stage. A direct relationship is then shown to exist between the DQE and values in the QAD. The QAD of a hypothetical system consisting of a charge-coupled device camera and a scintillating screen is evaluated as an illustrative example. The conventional zero-frequency analysis suggests two quantum sinks occur with approximately equal importance: one in the number of x rays, and one in the number of optical quanta. The spatial-frequency dependent analysis, however, shows the optical quantum sink becomes severe and dominates at nonzero frequencies. The necessary increase in gain or optical numerical aperture required to prevent the optical quantum sink for spatial frequencies of interest is determined from the QAD analysis. The visual impact of this nonzero spatial-frequency quantum sink is shown in images generated using a Monte Carlo simulation of the cascading process.

Algorithms

Effects of blur and eccentricity on differential spatial displacement discrimination.

Differential spatial displacement discrimination thresholds were determined for stimuli consisting of blobs with Gaussian spatial and temporal contrast envelopes. The stimuli were presented at detection threshold luminance contrast. The tasks were similar to the two-point discrimination acuity task and the three-dot alignment hyperacuity task. Thresholds were determined as a function of eccentricity along the horizontal meridian of the visual field (from 45 degrees nasal to 65 degrees temporal). The spatial spread or blur parameter of the blobs was adopted as a scale parameter. The results show that the performance of the visual system in differential spatial displacement discrimination tasks becomes progressively more homogeneous for a progressive increase in the blur parameter of the stimuli. Scaling (i) the three-blob alignment results with estimates of the cortical magnification factor and (ii) the two-blob separation discrimination results with their corresponding neural blur parameter shows an impressive isotropy and blur scale-invariance for the mechanisms mediating differential spatial displacement discrimination across the visual field. These results are interpreted in terms of a scaled sampling lattice model of the visual system, in combination with an automatic scale-selection mechanism.

Adult