Spatial Summation Research Articles

Temporal and spatial summation of laser heat stimuli in cultured nociceptive neurons of the rat

We studied the efficacy of a near-infrared laser (1475 nm) to activate rat dorsal root ganglion (DRG) neurons with short punctate radiant heat pulses (55 µm diameter) and investigated temporal and spatial summation properties for the transduction process for noxious heat at a subcellular level. Strength-duration curves (10–80 ms range) indicated a minimum power of 30.2mW for the induction of laser-induced calcium transients and a chronaxia of 13.9 ms. However, threshold energy increased with increasing stimulus duration suggesting substantial radial cooling of the laser spot. Increasing stimulus duration demonstrated suprathreshold intensity coding of calcium transients with less than linear gains (Stevens exponents 0.29/35mW, 0.38/60mW, 0.46/70mW). The competitive TRPV1 antagonist capsazepine blocked responses to short near-threshold stimuli and significantly reduced responses to longer duration suprathreshold heat. Heating 1/3 of the soma of a neuron was sufficient to induce calcium transients significantly above baseline (p < 0.05), but maximum amplitude was only achieved by centering the laser over the entire neuron. Heat-induced calcium increase was highest in heated cell parts but rapidly reached unstimulated areas reminiscent of spreading depolarization and opening of voltage-gated calcium channels. Full intracellular equilibrium took about 3 s, consistent with a diffusion process. In summary, we investigated transduction mechanisms for noxious laser heat pulses in native sensory neurons at milliseconds temporal and subcellular spatial resolution and characterized strength duration properties, intensity coding, and spatial summation within single neurons. Thermal excitation of parts of a nociceptor spread via both membrane depolarization and intracellular calcium diffusion.

A behavioral receptive field for ocular following in monkeys: Spatial summation and its spatial frequency tuning.

In human and non-human primates, reflexive tracking eye movements can be initiated at very short latency in response to a rapid shift of the image. Previous studies in humans have shown that only a part of the central visual field is optimal for driving ocular following responses. Herein, we have investigated spatial summation of motion information across a wide range of spatial frequencies and speeds of drifting gratings by recording short-latency ocular following responses in macaque monkeys. We show that optimal stimulus size for driving ocular responses cover a small (<20° diameter), central part of the visual field that shrinks with higher spatial frequency. This signature of linear motion integration remains invariant with speed and temporal frequency. For low and medium spatial frequencies, we found a strong suppressive influence from surround motion, evidenced by a decrease of response amplitude for stimulus sizes larger than optimal. Such suppression disappears with gratings at high frequencies. The contribution of peripheral motion was investigated by presenting grating annuli of increasing eccentricity. We observed an exponential decay of response amplitude with grating eccentricity, the decrease being faster for higher spatial frequencies. Weaker surround suppression can thus be explained by sparser eccentric inputs at high frequencies. A Difference-of-Gaussians model best renders the antagonistic contributions of peripheral and central motions. Its best-fit parameters coincide with several, well-known spatial properties of area MT neuronal populations. These results describe the mechanism by which central motion information is automatically integrated in a context-dependent manner to drive ocular responses.Significance statementOcular following is driven by visual motion at ultra-short latency in both humans and monkeys. Its dynamics reflect the properties of low-level motion integration. Here, we show that a strong center-surround suppression mechanism modulates initial eye velocity. Its spatial properties are dependent upon visual inputs' spatial frequency but are insensitive to either its temporal frequency or speed. These properties are best described with a Difference-of-Gaussian model of spatial integration. The model parameters reflect many spatial characteristics of motion sensitive neuronal populations in monkey area MT. Our results further outline the computational properties of the behavioral receptive field underpinning automatic, context-dependent motion integration.

Non-laboratory adaptation to study spatial summation of pain during COVID-19 pandemic

During COVID-19 pandemic associated with multiple restrictions research performance is significantly hampered. This experiment aimed to verify, whether it is possible to adapt the experimental pain induction technique and observe the spatial summation of pain (SSp) within the field-like environment (so that it is possible to conduct the experiment outside of the laboratory and obtain similar results). Two variants of the Cold Pressor Task (CPT) were chosen to induce the SSp effect: i) using a professional apparatus available in the laboratory and ii) with the use of a commercially available plastic box filled with ice and water and a thermometer to measure the temperature of the water. Volunteers’ (n = 9) hands were divided into 5 segments, which were then subsequently immersed in the cold water (5°C) for 60 seconds (5-minute intertrial interval). Each participant took part in two sessions, i.e., using laboratory equipment and its non-laboratory equivalent. Preliminary observations confirmed that it is possible to induce SSp using both laboratory devices and methods adapted to non-laboratory conditions. SSp effect size was found to be similar in both methods. Small differences in the level of reported pain were observed, but these differences did not affect the SSp effect. The study showed that it is possible to adapt the procedure to induce pain and SSp within non-laboratory environment.

Spatial Tuning in Nociceptive Processing Is Driven by Attention

When the source of nociception expands, one can observe an increase in pain. This effect is called spatial summation of pain (SSP) and has been a subject of several investigations. However, still, mechanisms shaping magnitude of this effect are largely unknown. The purpose of this study was to investigate how spatial attention modulates SSP, and thus reported pain. Forty pain-free volunteers (N=40, 20 females) participated in this within-subject experiment. Following training phase, volunteers took part in two sessions based on three different hand immersions in the cold water (Cold Pressor Task) with constant noxious temperature. Participants were asked to either immerse the ulnar side (A), radial side (B) or both sides (A+B, whole hand) and provide overall pain ratings they experienced. In case of immersions of merged sides (A+B) they were also asked to provide divided-attention rating(s), i.e., first pain in A then in B (or B then A) and directed- attention ratings (pain only in A or in B). Results confirmed clear SSP effect: Reported pain during immersions of A or B was less intense than pain during immersions of A+B (p < 0.001). Data also confirmed that spatial tuning was altered as SSP was fully abolished when participants provided two ratings in divided fashion (p < 0.001). Pain was significantly lowered when attention was directed only to one side (A or B) during immersion of A+B (p < 0.001). We conclude that SSP is attentionally driven phenomenon, and thus mechanisms of this effect must have a central component. SSP can be fully abolished when pain is rated in divided fashion or when attention is attracted to smaller painful area.

A Model of Pattern Separation by Single Neurons.

For efficient processing, spatiotemporal spike patterns representing similar input must be able to transform into a less similar output. A new computational model with physiologically plausible parameters shows how the neuronal process referred to as “pattern separation” can be very well achieved by single neurons if the temporal qualities of the output patterns are considered. Spike patterns generated by a varying number of neurons firing with fixed different frequencies within a gamma range are used as input. The temporal and spatial summation of dendritic input combined with theta-oscillating excitability in the output neuron by subthreshold membrane potential oscillations (SMOs) lead to high temporal separation by different delays of output spikes of similar input patterns. A Winner Takes All (WTA) mechanism with backward inhibition suffices to transform the spatial overlap of input patterns to much less temporal overlap of the output patterns. The conversion of spatial patterns input into an output with differently delayed spikes enables high separation effects. Incomplete random connectivity spreads the times up to the first spike across a spatially expanded ensemble of output neurons. With the expansion, random connectivity becomes the spatial distribution mechanism of temporal features. Additionally, a “synfire chain” circuit is proposed to reconvert temporal differences into spatial ones.

Spinal Nociception is Facilitated during Cognitive Distraction

The nociceptive withdrawal reflex (NWR) is a behavioral response to protect the body from noxious stimuli. The spatial characteristics of the stimulus modulate the reflex response to prevent damage to the affected tissue. Interneurons in the deep dorsal horn in the spinal cord encode the relationship between stimulus characteristics and the magnitude of the NWR and are also likely integrating spatial information of the nociceptive stimulus. The aim of this study was to use the NWR to investigate whether the spinal spatial integration of a simultaneous stimulus is modulated by shifting the attention of the participant towards (attention) or away from (distraction) the stimulus. We hypothesized that the descending activity shapes the receptive fields of the spinal neurons encoding spatial integration of nociception. Twenty healthy volunteers participated in the study. Single and simultaneous stimuli were delivered through two stimulating electrodes located in the arch and on the lateral side in the sole of the foot. The NWR was quantified by electromyography from the Tibialis Anterior and Biceps Femoris muscles during baseline and active tasks (attention and distraction). During the baseline task, spatial summation of the NWR was evoked during simultaneous stimulation. During the distraction task, the NWR was significantly larger compared to baseline, regardless of the sites being stimulated (single and simultaneous stimuli). In contrast, the NWR recorded during the attention task did not differ from baseline. These results further support that the spinal NWR pathway is under descending control which can be modulated by cognitive processes. The NWRs recorded over both proximal and distal muscles were similarly affected by the tasks, suggesting that the descending control affects the lower leg spinal system, with no discrimination between spinal segments.

Natural biomaterial honey-based resistive switching device for artificial synapse in neuromorphic systems

Resistive switching is a promising technology for artificial synapses, the most critical component and building block of a neural network for brain-inspired neuromorphic computing. The artificial synapse is capable of emulating a signal process and memory functions of biological synapses. The artificial synapse fabricated by natural bioorganic materials is essential for developing soft, flexible, and biocompatible electronics and sustainable, biodegradable, and environmentally friendly neuromorphic systems. In this work, a natural biomaterial—honey based resistive switching device—was demonstrated to emulate some important functionalities of biological synapses, including synaptic potentiation and depression, short-term and long-term memory, spatial summation, and shunting inhibition. The results indicate the potential of honey based resistive switching for artificial synaptic devices in renewable neuromorphic systems and bioelectronics.

Hierarchical timescales in the neocortex: Mathematical mechanism and biological insights

A cardinal feature of the neocortex is the progressive increase of the spatial receptive fields along the cortical hierarchy. Recently, theoretical and experimental findings have shown that the temporal response windows also gradually enlarge, so that early sensory neural circuits operate on short timescales whereas higher-association areas are capable of integrating information over a long period of time. While an increased receptive field is accounted for by spatial summation of inputs from neurons in an upstream area, the emergence of timescale hierarchy cannot be readily explained, especially given the dense interareal cortical connectivity known in the modern connectome. To uncover the required neurobiological properties, we carried out a rigorous analysis of an anatomically based large-scale cortex model of macaque monkeys. Using a perturbation method, we show that the segregation of disparate timescales is defined in terms of the localization of eigenvectors of the connectivity matrix, which depends on three circuit properties: 1) a macroscopic gradient of synaptic excitation, 2) distinct electrophysiological properties between excitatory and inhibitory neuronal populations, and 3) a detailed balance between long-range excitatory inputs and local inhibitory inputs for each area-to-area pathway. Our work thus provides a quantitative understanding of the mechanism underlying the emergence of timescale hierarchy in large-scale primate cortical networks.

Input from torus longitudinalis drives binocularity and spatial summation in zebrafish optic tectum

BackgroundA continued effort in neuroscience aims to understand the way brain circuits consisting of diverse neuronal types generate complex behavior following sensory input. A common feature of vertebrate visual systems is that lower-order and higher-order visual areas are reciprocally connected. Feedforward projections confer visual responsiveness to higher-order visual neurons while feedback projections likely serve to modulate responses of lower-order visual neurons in a context-dependent manner. Optic tectum is the largest first-order visual brain area in zebrafish and is reciprocally connected with the torus longitudinalis (TL), a second-order visual brain area that does not receive retinal input. A functional role for feedback projections from TL to tectum has not been identified. Here we aim to understand how this feedback contributes to visual processing.ResultsIn this study, we demonstrate that TL feedback projections to tectum drive binocular integration and spatial summation in a defined tectal circuit. We performed genetically targeted, cell type-specific functional imaging in tectal pyramidal neurons (PyrNs) and their two input neuron populations: retinal ganglion cells (RGCs) and neurons in TL. We find that PyrNs encode gradual changes in scene luminance using a complement of three distinct response classes that encode different light intensity ranges. Functional imaging of RGC inputs to tectum suggest that these response classes originate in the retina and RGC input specifies PyrN functional classes. In contrast, TL input serves to endow PyrNs with large, compound receptive fields that span both retinal hemifields.ConclusionsThese findings reveal a novel role for the zebrafish TL in driving binocular integration and spatial summation in tectal PyrNs. The neural circuit we describe generates a population of tectal neurons with large receptive fields tailored for detecting changes in the visual scene.

Spatial and Temporal Summation Across Frames Drive the Perception of Complex Glass Patterns

Spatial and Temporal Summation Across Frames Drive the Perception of Complex Glass Patterns

Passive localization of multiple sources using joint RSS and AOA measurements in spectrum sharing system

In spectrum sharing systems, locating multiple radiation sources can efficiently find out the intruders, which protects the shared spectrum from malicious jamming or other unauthorized usage. Compared to single-source localization, simultaneously locating multiple sources is more challenging in practice since the association between measurement parameters and source nodes are not known. Moreover, the number of possible measurements-source associations increases exponentially with the number of sensor nodes. It is crucial to discriminate which measurements correspond to the same source before localization. In this work, we propose a centralized localization scheme to estimate the positions of multiple sources. Firstly, we develop two computationally light methods to handle the unknown RSS-AOA measurements-source association problem. One method utilizes linear coordinate conversion to compute the minimum spatial Euclidean distance summation of measurements. Another method exploits the long-short-term memory (LSTM) network to classify the measurement sequences. Then, we propose a weighted least squares (WLS) approach to obtain the closed-form estimation of the positions by linearizing the non-convex localization problem. Numerical results demonstrate that the proposed scheme could gain sufficient localization accuracy under adversarial scenarios where the sources are in close proximity and the measurement noise is strong.

The impact of water depth and speed on muscle fiber activation of healthy dogs walking in a water treadmill

BackgroundWater treadmills are frequently used in the rehabilitation of dogs, for example with the purpose of re-building muscular strength after surgery. However, little is known about how different water depths and velocities affect the muscular workload during aquatic locomotion. This study used acoustic myography to assess hind limb muscle fiber activation in 25 healthy large-breed dogs walking in a water treadmill. Acoustic myography sensors were attached to the skin over the vastus lateralis of the quadriceps and the biceps femoris muscles. The dogs walked at two velocities (30 and 50 m/min) and four water depths: bottom of the pads, hock, stifle and mid-femur. Acoustic myograph signals were analyzed for changes in three muscle function parameters: efficiency/coordination (E-score) and spatial (S-score) and temporal (T-score) summation.ResultsDifferences between E, S, and T were statistically significant compared across different speeds (30, 50) and water levels (0, 1, 2, 3) using a one-way ANOVA with multiple comparisons (Tukey; Geisser-Greenhouse correction) as well as a two-tailed one sample t-test. At 50 m/min in water at the mid-femur, the biceps femoris was less efficient (P < 0.001) and recruited more fibers (P = 0.01) at a higher firing rate (P = 0.03) compared to working in shallower water, while the vastus lateralis was also less efficient (P < 0.01), but spatial and temporal summation did not change significantly. At 30 m/min, biceps efficiency was reduced (P < 0.01) when water was at the mid-femur compared to the bottom of the pads level. Walking in stifle- or hock-deep water did not show increased muscle activation for either muscle compared to walking in water at the bottom of the pads.ConclusionMore muscle activation was required to walk in water at a depth at the level of the mid-femur compared to shallower water, and this exercise was more demanding for the biceps femoris, a muscle engaged in propulsion, than for vastus lateralis. These findings may help practitioners towards making more precise rehabilitation protocols.

Compartmental spiking neuron model for pattern classification

One of the directions of development within the framework of the neuromorphic approach is the development of anatomically similar models of brain networks, taking into account the structurally complex structure of neurons and the adaptation of connections between them, as well as the development of learning algorithms for such models. In this work, we use the previously presented compartmental spike model of a neuron, which describes the structure (dendritic tree, soma, synapses) and behaviour (temporal and spatial signal summation, generation of action potential, stimulation and suppression of electrical activity) of a biological neuron. An algorithm for the structural organization of neuron models into a spike neural network is proposed for recognizing an arbitrary impulse pattern by introducing inhibitory synapses between trained neuron models. The dynamically adapting neuron models used are trained according to a previously proposed algorithm that automatically selects parameters such as soma size, dendrite length, and the number of synapses on each of the dendrites in order to induce a temporal response at the output depending on the input pattern encoded using a time window and temporal delays in the vector of single spikes arriving at a separate dendrite of a neuron. The developed algorithms are evaluated on the Iris dataset classification problem with four training examples from each class. As a result of the classification, separate disjoint clusters are formed, which demonstrates the applicability of the proposed spike neural network with a dynamically changing structure of elements in the problem of pattern recognition and classification.

Synthesis of a comprehensive population code for contextual features in the awake sensory cortex.

How cortical circuits build representations of complex objects is poorly understood. Individual neurons must integrate broadly over space, yet simultaneously obtain sharp tuning to specific global stimulus features. Groups of neurons identifying different global features must then assemble into a population that forms a comprehensive code for these global stimulus properties. Although the logic for how single neurons summate over their spatial inputs has been well explored in anesthetized animals, how large groups of neurons compose a flexible population code of higher-order features in awake animals is not known. To address this question, we probed the integration and population coding of higher-order stimuli in the somatosensory and visual cortices of awake mice using two-photon calcium imaging across cortical layers. We developed a novel tactile stimulator that allowed the precise measurement of spatial summation even in actively whisking mice. Using this system, we found a sparse but comprehensive population code for higher-order tactile features that depends on a heterogeneous and neuron-specific logic of spatial summation beyond the receptive field. Different somatosensory cortical neurons summed specific combinations of sensory inputs supra-linearly, but integrated other inputs sub-linearly, leading to selective responses to higher-order features. Visual cortical populations employed a nearly identical scheme to generate a comprehensive population code for contextual stimuli. These results suggest that a heterogeneous logic of input-specific supra-linear summation may represent a widespread cortical mechanism for the synthesis of sparse higher-order feature codes in neural populations. This may explain how the brain exploits the thalamocortical expansion of dimensionality to encode arbitrary complex features of sensory stimuli.

Spatial Acuity of the Nociceptive System and Spatial Summation of Pain: Potential Implications for the Clinic

Previous studies investigating spatial acuity measured by two-point discrimination threshold concluded that the nociceptive system is less accurate than the innocuous tactile system. In the discussed article, the authors point out that the nociceptive system is more accurate than the tactile system when controlling for the stimulus modality and intensity in healthy pain-free individuals. Furthermore, this article shows that the pattern of distance-based and areabased spatial summation of pain is modality independent.

Optical Sensitivity of Camera-Like Eyes to White Light.

sGastropod mollusks are convenient model organisms for studying the functioning of the visual system. The purpose of this work is to estimate the value of the optical sensitivity to white light for the camera-like eyes of gastropod mollusks and humans and analyze its effect on the spatial resolving power in two regions of the retina: in the center—for single photoreceptors of the first/second type in a mollusk and single cones in humans—and in the periphery—for single photoreceptors of the first/second type in a mollusk, as well as for single rods/cones and their groups, subject to spatial summation in humans. The methods of histology, light and transmission electron microscopy, morphometry, calculations and methods of statistical analysis are used in the work. In a mollusk, with a fixed pupil area, the value of the optical sensitivity of the eye to white light in the center of the retina for single photoreceptors of the first/second type is 0.5/0.006 μm2·sr and in the periphery of the retina, 0.9/0.009 μm2·sr. In humans, at the minimum and maximum pupil area, respectively, the value of the optical sensitivity of the eye to white light in the center of the retina (foveola) for single cones varies from 0.00053 to 0.028 μm2·sr, and in the periphery of the retina (far periphery) for single rods from 0.011 to 0.575 μm2·sr, for single cones from 0.025 to 1.319 μm2·sr, for the groups of rods from 3859 to 204,094 μm2·sr and for the groups of cones from 2.5 to 131 μm2·sr. The value of the optical sensitivity of the eyes to white light for single photoreceptors of the first/second type in both regions of the retina in a mollusk, as well as for single cones in the center and groups of rods/cones in the periphery of the retina in humans, corresponds to the ambient light conditions during periods of activity and does not affect the spatial resolving power.

Spatial suppression of chromatic motion

Center-surround antagonism, as a ubiquitous feature in visual processing, usually leads to inferior perception for a large stimulus compared to a small one. For example, it is more difficult to judge the motion direction of a large high-contrast pattern than that of a small one. However, this spatial suppression in the motion dimension was only reported for luminance motion, and was not found for chromatic motion. Given that center-surround suppression only occurs for strong visual inputs, we hypothesized that previous failure in finding spatial suppression of chromatic motion might be due to weak chromatic motion being induced with stimuli of limited parameters. In this study, we used phase-shift discrimination and motion-direction discrimination tasks to measure motion spatial suppression induced by stimuli of two spatial frequencies (0.5 and 2 cpd) and two contrasts (low and high). We found that spatial suppression of the chromatic motion was stably observed for stimuli of high spatial frequency (2 cpd) and high contrast and spatial summation occurred for stimuli of low spatial frequency (0.5 cpd). Intriguingly, there was no correlations between the motion spatial suppressions of luminance motion and chromatic motion, implying that the two types of spatial suppression are not originated from the same neural processing. Our findings indicate that spatial suppression also exists for chromatic motion, and the mechanisms underlying the spatial suppression of chromatic motion is different from that of luminance motion.

Robust prediction metrics for structure-borne noise in timber buildings

This paper presents an investigation of prediction metrics for structure-borne noise in timber buildings, based on the vibration response of the structure. The purpose of the study is to derive a robust measure for predicting structure-borne noise in timber buildings based solely on the vibration response; a measure with a known correlation to the actual acoustic response. Such a measure is useful for establishing simplified numerical prediction models, and for making robust assessments of different design alternatives. The procedure has previously been applied to automotive applications with a valuable outcome for the industry. The vibration and sound pressure responses in a room of a multi-storey building are calculated using the finite element method. As excitation, a harmonic unit point load (20-300 Hz) located at another storey of the building is used. Based on the structural and acoustic response fields, scalar values are calculated by performing spatial and frequency domain averaging and summations. Various methods for calculating scalar values from the vibration response are investigated. The correlation between the vibration scalar values and the acoustic scalar values are presented.

Selective postsaccadic enhancement of motion perception

Saccadic eye movements can drastically affect motion perception: during saccades, the stationary surround is swept rapidly across the retina and contrast sensitivity is suppressed. However, after saccades, contrast sensitivity is enhanced for color and high-spatial frequency stimuli and reflexive tracking movements known as ocular following responses (OFR) are enhanced in response to large field motion. Additionally, OFR and postsaccadic enhancement of neural activity in primate motion processing areas are well correlated. It is not yet known how this postsaccadic enhancement arises. Therefore, we tested if the enhancement can be explained by changes in the balance of centre-surround antagonism in motion processing, where spatial summation is favoured at low contrasts and surround suppression is favoured at high contrasts. We found motion perception was selectively enhanced immediately after saccades for high spatial frequency stimuli, consistent with previously reported selective postsaccadic enhancement of contrast sensitivity for flashed high spatial frequency stimuli. The observed enhancement was also associated with changes in spatial summation and suppression, as well as contrast facilitation and inhibition, suggesting that motion processing is augmented to maximise visual perception immediately after saccades. The results highlight that spatial and contrast properties of underlying neural mechanisms for motion processing can be affected by an antecedent saccade for highly detailed stimuli and are in line with studies that show behavioural and neuronal enhancement of motion processing in non-human primates.

Vibration Perception Thresholds of Skin Mechanoreceptors Are Influenced by Different Contact Forces.

Determining vibration perception thresholds (VPT) is a central concern of clinical research and science to assess the somatosensory capacity of humans. The response of different mechanoreceptors to an increasing contact force has rarely been studied. We hypothesize that increasing contact force leads to a decrease in VPTs of fast-adapting mechanoreceptors in the sole of the human foot. VPTs of 10 healthy subjects were measured at 30 Hz and 200 Hz at the heel of the right foot using a vibration exciter. Contact forces were adjusted precisely between 0.3 N–9.6 N through an integrated force sensor. Significant main effects were found for frequency and contact force. Furthermore, there was a significant interaction for frequency and contact force, meaning that the influence of an increasing contact force was more obvious for the 30 Hz condition. We presume that the principles of contrast enhancement and spatial summation are valid in Meissner and Pacinian corpuscles, respectively. In addition to spatial summation, we presume an effect on Pacinian corpuscles due to their presence in the periosteum or interosseous membrane.