Showing papers on "Summation published in 2004"

Natural stimulus statistics alter the receptive field structure of v1 neurons.

Abstract: Studies of the primary visual cortex (V1) have produced models that account for neuronal responses to synthetic stimuli such as sinusoidal gratings. Little is known about how these models generalize to activity during natural vision. We recorded neural responses in area V1 of awake macaques to a stimulus with natural spatiotemporal statistics and to a dynamic grating sequence stimulus. We fit nonlinear receptive field models using each of these data sets and compared how well they predicted time-varying responses to a novel natural visual stimulus. On average, the model fit using the natural stimulus predicted natural visual responses more than twice as accurately as the model fit to the synthetic stimulus. The natural vision model produced better predictions in >75% of the neurons studied. This large difference in predictive power suggests that natural spatiotemporal stimulus statistics activate nonlinear response properties in a different manner than the grating stimulus. To characterize this modulation, we compared the temporal and spatial response properties of the model fits. During natural stimulation, temporal responses often showed a stronger late inhibitory component, indicating an effect of nonlinear temporal summation during natural vision. In addition, spatial tuning underwent complex shifts, primarily in the inhibitory, rather than excitatory, elements of the response profile. These differences in late and spatially tuned inhibition accounted fully for the difference in predictive power between the two models. Both the spatial and temporal statistics of the natural stimulus contributed to the modulatory effects.

Effects of diffuse noxious inhibitory controls on temporal summation of the RIII reflex in humans.

Abstract: The aim of this study was to investigate the effects of diffuse noxious inhibitory controls (DNICs) on the temporal summation of the nociceptive flexion reflex (RIII reflex) in humans. Recordings were obtained from 36 healthy adults (16 M, 20 F), and the area and temporal summation threshold (TST) of the RIII reflex were measured. The subjective intensity of the painful sensation was rated on an 11-point visual analogue scale (VAS). Neurophysiological and VAS measurements were recorded after activation of DNICs by means of the cold pressor test (CPT), which involved immersing the hand in cold water (2-4 degrees C). A slight significant lower TST was found in the females versus the males. In all the subjects, the CPT induced a significant TST increase and RIII area reduction compared with the control session. The VAS results paralleled those of the RIII reflex area and TST. During the CPT, a significant difference in the percentage TST increase emerged between females and males, being lower in the former. Similarly, we found a significantly lower percentage reduction of the RIII area in women than in men during the CPT. To summarize, activation of DNICs through the CPT significantly increased the TST of the RIII reflex in healthy subjects. This inhibitory effect was gender-specific. Whereas other findings are based on psychophysical evaluations, the results of this experimental study provide an objective neurophysiological demonstration that DNICs attenuate temporal summation in humans and confirm the presence of significant differences in pain modulation mechanisms between men and women.

Sex differences in temporal summation of pain and aftersensations following repetitive noxious mechanical stimulation

Abstract: Several studies demonstrate that women are more sensitive to experimental pain than men. In addition, women exhibit greater temporal summation of heat and mechanically evoked pain. Since temporal summation of pain is centrally mediated, its greater expression in women suggests a central nociceptive hyperexcitability relative to men. The purpose of this study was to pursue this theory, by further assessing sex differences in (1) temporal summation of mechanically evoked pain, and (2) aftersensations following repetitive noxious stimulation. Sixteen series of 10 repetitive, mildly noxious, mechanical stimuli were applied to the fingers of 25 women and 25 age-matched men. The subjects rated the pain intensity and unpleasantness caused by the first, fifth and tenth stimulus in the series, as well as their aftersensations 15 s and 1 min following the end of stimulation. Data were analyzed by three-way repeated-measures analysis of variance. Pain and unpleasantness ratings increased with repetition of stimulation (P<0.0001). Temporal summation of pain intensity and unpleasantness ratings were more pronounced in women than men (P<0.0001). In addition, significant temporal summation occurred only with 2 s interstimulus interval for men (P<0.0005) but with 2 and 5 s interstimulus interval for women (P<0.0001). Moreover, women provided greater ratings for the intensity and the unpleasantness of aftersensations (P<0.0005) and reported painful aftersensations at greater frequency (P<0.05) Greater temporal summation of pain and aftersensations in women suggests that their central processing of nociceptive input may be more easily upregulated into pathological hyperexcitability, possibly accounting for the higher prevalence of various chronic pain conditions among women.

Dendritic Ih selectively blocks temporal summation of unsynchronized distal inputs in CA1 pyramidal neurons.

Abstract: The active dendritic conductances shape the input-output properties of many principal neurons in different brain regions, and the various ways in which they regulate neuronal excitability need to be investigated to better understand their functional consequences. Using a realistic model of a hippocampal CA1 pyramidal neuron, we show a major role for the hyperpolarization-activated current, Ih, in regulating the spike probability of a neuron when independent synaptic inputs are activated with different degrees of synchronization and at different distances from the soma. The results allowed us to make the experimentally testable prediction that the Ih in these neurons is needed to reduce neuronal excitability selectively for distal unsynchronized, but not for synchronized, inputs.

Spatio-temporal luminance contrast sensitivity and visual backward masking in schizophrenia

Abstract: The aim of two experiments was to investigate the relationship between spatio-temporal contrast sensitivity and visual backward masking in normal observers and in subgroups with positive or negative symptoms in schizophrenia. Experiment 1 measured contrast sensitivity for stationary and counterphase-modulated sinusoidal gratings at four spatial (0.5, 2.0, 4.0, 8.0 cycles/degree) and four temporal frequencies (0, 4.0, 8.0, 16.0 Hz). The results showed that there were no differences in spatio-temporal contrast sensitivity between the control and positive-symptom group, and in comparison with these groups, contrast sensitivity was significantly lower at all spatial and temporal frequencies in the negative-symptom group. Experiment 2 measured the visibility of a Landolt C target with a constant target stimulus duration of 4.0 ms followed by a 150-ms backward mask, which was presented at 12 stimulus onset asynchronies from 0 to 110 ms in the same groups of observers. Consistent with the findings of the previous experiment, there were no significant differences in backward masking between the control and positive-symptom group, and in comparison with these groups, visual backward masking was significantly higher at all stimulus onset asynchronies from 40 to 110 ms in the negative-symptom group. The present findings show that there were no significant differences in contrast sensitivity and in backward masking between normal observers and a group with positive symptoms in schizophrenia. It was concluded that the reduction in contrast sensitivity for low spatial frequency counterphase flicker in the negative-symptom group is consistent with a reduction in the 'contrast gain control' mechanism of magnocellular channels, and that the reduction in contrast sensitivity for medium and high stationary gratings is consistent with a disorder in parvocellular channels. It was proposed that a disorder in magnocellular channels in the negative-symptom group may enforce a reliance on parvocellular channels that results in longer temporal summation and visible persistence, slower visual processing of single target stimuli at threshold and higher levels of sensory integration, and backward masking when multiple stimuli are presented rapidly in time.

Spatial summation of heat pain within and across dermatomes in fibromyalgia patients and pain-free subjects.

Abstract: The mechanisms of spatial summation of pain (SSP) include pain coding dependent on impulse frequency and the number of recruited central neurons. However, SSP may also be influenced by pain inhibitory mechanisms, such as diffuse noxious inhibitory controls. Abnormal interactions between pain inhibitory mechanisms and SSP may be relevant for chronic pain conditions such as fibromyalgia (FM) and may help explain why widespread pain is characteristic for this chronic pain syndrome. The present study was designed to determine the difference of thermal SSP in the upper extremities between FM and normal control (NC) subjects, particularly within and across dermatomes of the hand. Fourteen NC and 19 FM subjects were enrolled in this study. SSP testing sessions involved immersion of each individual fingertip as well as stepwise immersion of the fingers, hands, and forearms in a hot water bath (46 °C) for 5s and 20s. In addition, immersion of several fingertips across dermatome C 7 –C 8 was compared to progressive immersion of the index finger (dermatome C 7 ). These experiments demonstrated significant spatial summation of heat-induced pain in both FM and NC subjects. SSP was most extensive within the fingers, and became negligible as the stimulus area increased above the hand. Furthermore, SSP was more pronounced within one dermatome such as that of the index finger than across several dermatomes of the hand. These results were similar for both FM and NC subjects. Thus, mechanisms of SSP, including possible inhibitory factors that limit this relevant pain mechanism, appear to be similar for both FM and NC subjects.

Contrast-Dependent Spatial Summation in the Lateral Geniculate Nucleus and Retina of the Cat

Abstract: Based on extracellular recordings from 69 lateral geniculate nucleus (LGN) cells in the anesthetized cat, we found spatial summation within their receptive fields to be dependent on the contrast of the stimuli presented. By fitting the summation curves to a difference of Gaussians model, we attributed this contrast-dependent effect to an actual change in the size of the center mechanism. Analogous changes in spatial frequency tuning were also observed, specifically increased peaks and cut-off frequencies with contrast. These effects were seen across the populations of both X and Y cell types. In a few cases, LGN cells were recorded simultaneously with one of their retinal ganglion cell (RGC) inputs (S-potentials). In every case, the RGCs exhibited similar contrast-dependent effects in the space and spatial-frequency domains. We propose that this contrast dependency in the retinal ganglion cells results directly from a reduction in the size of the center mechanism due to an increase in contrast. We also propose that these properties first arise in the retina and are transmitted passively through the LGN to visual cortex.

Synaptic efficacy during repetitive activation of excitatory inputs in primate dorsolateral prefrontal cortex.

Abstract: Neurons in the monkey dorsolateral prefrontal cortex (DLPFC) fire persistently during the delay period of working memory tasks. To determine how repetitive firing affects the efficacy of synaptic inputs to DLPFC layer 3 neurons, we examined the effects of repetitive presynaptic stimulation on the amplitude and temporal summation of EPSPs. Recordings were obtained in monkey DLPFC brain slices from regular spiking (RS) pyramidal cells and two types of interneurons, fast spiking (FS) and adapting non-pyramidal (ANP) cells. Repetitive stimulation of presynaptic axons in layer 3 caused EPSP depression in RS and FS neurons, but EPSP facilitation in ANP cells. A shorter EPSP duration produced weaker temporal summation in FS neurons compared to the other cell classes. Thus, due to the combined effects of dynamic changes in EPSP amplitude and differences in temporal summation, the effect of a presynaptic spike train differed according to the postsynaptic cell class. Similar results were obtained when recording unitary EPSPs evoked in connected pairs of presynaptic RS pyramidal cells and postsynaptic RS, FS or ANP neurons. In addition, similar differences in the efficacy of sustained inputs among cell classes were observed when delay-related firing was reproduced in vitro by stimulating inputs with the timing of spike trains recorded from the DLPFC of monkeys performing a delayed-response task. We suggest that the transition from baseline firing rates to higher frequency delay-related firing may lead to the differential activation of distinct cell populations, with corresponding significant effects on the patterns of activity in local prefrontal circuits.

Investigation of the facilitation of the nociceptive withdrawal reflex evoked by repeated transcutaneous electrical stimulations as a measure of temporal summation in conscious horses.

Abstract: Objective—To investigate whether facilitation of the nociceptive withdrawal reflex (NWR) can be evoked and quantified as a measure of temporal summation from the distal aspect of the left forelimb and hind limb in standing nonsedated horses via repeated stimulations of various subthreshold intensities and frequencies. Animals—10 adult horses. Procedure—Surface electromyographic activity evoked by stimulation of the digital palmar and plantar nerves was recorded from the common digital extensor and cranial tibial muscles. For each horse, the NWR threshold intensity to a single stimulus was determined for the forelimb and hind limb. Repeated stimulations were performed at subthreshold intensities and at frequencies of 2, 5, and 10 Hz. The reflex amplitude was quantified, and the behavioral responses accompanying the stimulations were scored. Results—Repeated stimulations at subthreshold intensities were able to summate and facilitate the NWR in conscious horses. The reflex facilitation was significantly rel...

Contrast dependence of spatial summation revealed by classification image analysis

Abstract: Detection of low-contrast luminance-defined stimuli can involve spatial summation over a large portion of the visual field. However prior psychophysical results suggest that the summation region may shrink substantially in the presence of high-contrast masking gratings or noise (Legge & Foley, 1980; Kersten, 1984). This may be related to recent findings that the extent of spatial summation in V1 neurons depends upon contrast (Sceniak et al., 1999; Kapadia et al., 1999). Here we use a classification image technique to directly test whether the psychophysical receptive field for a simple stimulus (a vertical edge in noise) is dependent upon contrast.

The cortically evoked secondary depolarization affects the integrative properties of thalamic reticular neurons.

Abstract: Corticothalamic terminals on thalamic reticular (RE) neurons account for most synapses from afferent pathways onto this nucleus and these inputs are more powerful than those from axon collaterals of thalamocortical neurons. Given the supremacy of cortical inputs, we analysed here the characteristics and possible mechanisms underlying a secondary component of the cortically elicited depolarization in RE neurons, recorded in cats under barbiturate anesthesia. Electrical stimulation of corticothalamic axons in the internal capsule evoked fixed and short-latency excitatory postsynaptic potentials (EPSPs) that, by increasing stimulation intensity and at hyperpolarized levels (< -70 mV), developed into low-threshold spikes and spindle oscillations. The threshold for spindle oscillations was 60% higher than that required for evoking minimal EPSPs. The evoked EPSPs included a secondary depolarizing component, which appeared approximately 5 ms after the peak of the initial component and was voltage dependent, i.e. most prominent between -70 mV and -85 mV, while being greatly reduced or absent at more hyperpolarized levels. The secondary depolarizing component was sensitive to QX-314 in the recording micropipette. We suggest that the secondary component of cortically evoked EPSPs in RE neurons is due to the dendritic activation of T-currents, with a probable contribution of the persistent Na+ current. This late component affected the integrative properties of RE neurons, including their spiking output and temporal summation of incoming cortical inputs.

A single mechanism for both luminance and chromatic grating vernier tasks: evidence from temporal summation.

Abstract: Vernier thresholds are determined by luminance rather than chromatic contrast when both are present in vernier targets. The role of luminance and chromatic mechanisms in vernier performance under equiluminant conditions remains uncertain. Temporal summation functions for vernier thresholds with luminance and red-green equiluminant gratings were compared to those for detection thresholds with similar stimuli. Vernier thresholds showed similar temporal summation for luminance and chromatic gratings, which is consistent with a single mechanism underlying vernier performance in the two conditions. However, detection thresholds showed a shorter temporal summation duration for luminance gratings than for chromatic gratings, which suggests that two different mechanisms underlie detection thresholds. Analysis of physiological data supports the hypothesis that the frequency-doubled response of ganglion cells in the magnocellular pathway can provide accurate spatiotemporal information for vernier performance at equiluminance.

Comparison of hyperalgesia induced by capsaicin injection and controlled heat injury: effect on temporal summation.

Abstract: The relationship between induction of central sensitization and facilitation of temporal summation to repetitive stimulation is still unclear. The aim of this study was to investigate temporal summation before and after the induction of secondary hyperalgesia by two different experimental methods: capsaicin injection and controlled heat injury. The effect of each injury model was assessed on a separate day with an interval of at least 5 days. Twelve healthy volunteers participated. Each experiment was performed using electrical, radiant heat, mechanical impact, and punctuate stimuli consecutively. The pain threshold (PT) to a single stimulus and the summation threshold to five repetitive stimuli for electrical (2 Hz) and radiant heat (0.83 Hz) were assessed within the secondary hyperalgesic area. The degree of temporal summation for stimulus intensities of 0.8, 1.0, and 1.2 times the baseline pain thresholds were evaluated by the increase in visual analogue scale (VAS) scores from the first to the fifth s...

CHAPTER 16 – Postsynaptic Potentials and Synaptic Integration

Abstract: This chapter discusses the postsynaptic potentials (PSPs) and synaptic integration. PSPs in the central nervous system can be divided into two broad classes on the basis of mechanisms and the duration of potential. One class is based on the direct binding of a transmitter molecule(s) with a receptor–channel complex; these receptors are ionotropic. The other class of PSPs is based on the indirect effect of transmitter molecule(s) binding with a receptor. The receptors that produce these PSPs are metabotropic. The sign of the postsynaptic potential is determined by the relationship between the membrane potential of the postsynaptic neuron and the ion selectivity of the ionotropic receptor. The low threshold Na + and Ca 2+ channels can boost an excitatory postsynaptic potential (EPSP). It is considered that the spatial distribution of the various voltage-dependent channels, ligand-gated receptors, and metabotropic receptors is not uniform. Thus, each segment of the neuronal membrane can perform selective integrative functions. The system has an enormous capacity for the local processing of information and for performing logical operations.