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Real-time visualization of neural synchrony for identifying coordinated cell assemblies.

We introduce a synchrony map that translates the fine temporal organization of multi-unit responses in the visual cortex into an easily interpreted spatial display. We test the synchrony map on microelectrode array recordings in Area 17 of anesthetized and paralyzed cats. We first examine the synchrony map using averaged data and probability calculations to demonstrate orientation-dependent changes in synchrony. We then demonstrate how the synchrony map can be implemented for real-time visualization of synchrony among neural assemblies.

Action Potentials↗

Cross-modal perceptual integration of spatially and temporally disparate auditory and visual stimuli.

Under certain conditions, auditory and visual information are integrated into a single unified percept even when they originate in different locations in space. The present study shows how this illusion, known as the ventriloquism effect, depends on spatial, temporal and cognitive factors. A method of psychophysical scaling was employed in combination with simple auditory-visual stimuli (tone bursts and flashing light spots) that were presented with various spatiotemporal disparities. Participants either judged their impression of the likelihood of a common cause (Experiment 1) or spatial alignment (Experiment 2) or synchrony of sound and light (Experiment 3). In all three experiments the participants' judgements depended significantly on temporal disparity whereas influences of spatial disparity were significant in Experiments 1 and 2. Optimum scores were always obtained when auditory stimuli were presented with a delay of 50-100 ms after the visual stimuli. These results demonstrate that both temporal and spatial proximity of the two stimuli are critical for the experience of phenomenal causality. On the other hand, spatio-temporal ranges for optimal perception of phenomenal causality in Experiment 1 were significantly larger than predicted by simultaneous detection of spatial and temporal disparities. This finding suggests that auditory-visual binding was further facilitated by additional, cognitive, factors, associated with the specific instruction to judge the likelihood of a common cause. Obviously, these instructional influences may reflect similar perceptual effects, as have been shown previously by increasing the complexity or cognitive compellingness of auditory-visual stimuli.

Acoustic Stimulation↗

Phenology of forest caterpillars and their host trees: the importance of synchrony.

For many leaf-feeding herbivores, synchrony in phenology with their host plant is crucial as development outside a narrow phenological time window has severe fitness consequences. In this review, we link mechanisms, adaptation, and population dynamics within a single conceptual framework, needed for a full understanding of the causes and consequences of this synchrony. The physiological mechanisms underlying herbivore and plant phenology are affected by environmental cues, such as photoperiod and temperature, although not necessarily in the same way. That these different mechanisms lead to synchrony, even if there is spatial and temporal variation in plant phenology, is a result of the strong natural selection acting on the mechanism underlying herbivore phenology. Synchrony has a major impact on the population densities of leaf-feeding Lepidoptera, and years with a high synchrony may lead to outbreaks. Global climate change leads to a disruption of the synchrony between herbivores and their host plants, which may have major impacts for population viability if natural selection is insufficient to restore synchrony.

Animals↗

Spatial summation in the receptive fields of simple cells in the cat's striate cortex.

1. We have examined the responses of simple cells in the cat's atriate cortex to visual patterns that were designed to reveal the extent to which these cells may be considered to sum light-evoked influences linearly across their receptive fields. We used one-dimensional luminance-modulated bars and grating as stimuli; their orientation was always the same as the preferred orientation of the neurone under study. The stimuli were presented on an oscilloscope screen by a digital computer, which also accumulated neuronal responses and controlled a randomized sequence of stimulus presentations. 2. The majority of simple cells respond to sinusoidal gratings that are moving or whose contrast is modulated in time in a manner consistent with the hypothesis that they have linear spatial summation. Their responses to moving gratings of all spatial frequencies are modulated in synchrony with the passage of the gratings' bars across their receptive fields, and they do not produce unmodulated responses even at the highest spatial frequencies. Many of these cells respond to temporally modulated stationary gratings simply by changing their response amplitude sinusoidally as the spatial phase of the grating the grating is varied. Nonetheless, their behavior appears to indicate linear spatial summation, since we show in an Appendix that the absence of a 'null' phase in a visual neurone need not indicate non-linear spatial summation, and further that a linear neurone lacking a 'null' phase should give responses of the form that we have observed in this type of simple cell. 3. A minority of simple cells appears to have significant non-linearities of spatial summation. These neurones respond to moving gratings of high spatial frequency with a partially or totally unmodulated elevation of firing rate. They have no 'null' phases when tested with stationary gratings, and reveal their non-linearity by giving responses to gratings of some spatial phases that are composed partly or wholly of even harmonics of the stimulus frequency ('on-off' responses). 4. We compared simple receptive fields with their sensitivity to sinusoidal gratings of different spatial frequencies. Qualitatively, the most sensitive subregions of simple cells' receptive fields are roughly the same width as the individual bars of the gratings to which they are most sensitive. Quantitatively, their receptive field profiles measured with thin stationary lines, agree well with predicted profiles derived by Fourier synthesis of their spatial frequency tuning curves.

Action Potentials↗

Global and local synchrony of coupled neurons in small-world networks.

Synchronous firing of neurons is thought to play important functional roles such as feature binding and switching of cognitive states. Although synchronization has mainly been investigated so far using model neurons with simple connection topology, real neural networks have more complex structures. Here we examine the behavior of pulse-coupled leaky integrate-and-fire neurons with various network structures. We first show that the dispersion of the number of connections for neurons influences dynamical behavior even if other major topological statistics are kept fixed. The rewiring probability parameter representing the randomness of networks bridges two spatially opposite frameworks: precise local synchrony and rough global synchrony. Finally, cooperation of the global connections and the local clustering property, which is prominent in small-world networks, forces synchrony of distant neuronal groups receiving coherent inputs.

Action Potentials↗

Synchrony & chaos in patchy ecosystems.

The apparent synchronisation of spatially discrete populations is a well documented phenomenon. However, it is not clear what the governing mechanisms are for this synchrony, and whether they are robust over a range of environmental conditions and patch specific population dynamic behaviours. In this paper, we explore two (possibly interacting) modes of coupling, and investigate their theoretically discernible, and perhaps even experimentally measurable, signatures. To aid us in this investigation we employ a planktonic example system, with direct application to plankton patchiness. Furthermore, we address the role of chaos in complex spatio-temporal dynamics; we find that chaos associated with funnel attractors can play a distinguished role, over dynamics less sensitive to small variations, in being more susceptible to generalised synchronisation (such as phase synchronisation) in the presence of small local parameter variation. This is in contrast to the case for coupled systems with identical dynamics, and suggests that non-identically coupled systems are more vulnerable to global extinction events when exhibiting funnel-type chaotic dynamics.

Animals↗

Ensemble patterns of hippocampal CA3-CA1 neurons during sharp wave-associated population events.

Transfer of neuronal patterns from the CA3 to CA1 region was studied by simultaneous recording of neuronal ensembles in the behaving rat. A nonlinear interaction among pyramidal neurons was observed during sharp wave (SPW)-related population bursts, with stronger synchrony associated with more widespread spatial coherence. SPW bursts emerged in the CA3a-b subregions and spread to CA3c before invading the CA1 area. Synchronous discharge of >10% of the CA3 within a 100 ms window was required to exert a detectable influence on CA1 pyramidal cells. Activity of some CA3 pyramidal neurons differentially predicted the ripple-related discharge of circumscribed groups of CA1 pyramidal cells. We suggest that, in SPW behavioral state, the coherent discharge of a small group of CA3 cells is the primary cause of spiking activity in CA1 pyramidal neurons.

Action Potentials↗

Simultaneous rate-synchrony codes in populations of spiking neurons.

Firing rates and synchronous firing are often simultaneously relevant signals, and they independently or cooperatively represent external sensory inputs, cognitive events, and environmental situations such as body position. However, how rates and synchrony comodulate and which aspects of inputs are effectively encoded, particularly in the presence of dynamical inputs, are unanswered questions. We examine theoretically how mixed information in dynamic mean input and noise input is represented by dynamic population firing rates and synchrony. In a subthreshold regime, amplitudes of spatially uncorrelated noise are encoded up to a fairly high input frequency, but this requires both rate and synchrony output channels. In a suprathreshold regime, means and common noise amplitudes can be simultaneously and separately encoded by rates and synchrony, respectively, but the input frequency for which this is possible has a lower limit.

Action Potentials↗

Closer in time when farther in space--spatial factors in audiovisual temporal integration.

We investigated the effect of visual eccentricity and spatial alignment on judgments of audiovisual synchrony. Sequences of flashes at 4, 6, and 8 Hz were presented centrally, or at horizontal eccentricities of 6 degrees or 18 degrees. Concurrent sequences of clicks were presented at the same rate as the flashes, or at higher or lower rates. Subjects judged whether the flash rate was the same as (synchronous with), faster than, or slower than the click rate. With the 4- and 6-Hz flash rates, subjects' judgments of audiovisual synchrony increased with increasing eccentricity, but only when the click rate was more rapid than the flash rate. This effect remained even when the size of the peripheral visual stimuli was adjusted to compensate for cortical magnification, and was not significantly influenced by the spatial proximity of the auditory and visual signals. However, it was absent when the auditory and visual stimuli were presented serially rather than concurrently. With the 8-Hz flash rate, synchrony judgments were prevalent irrespective of eccentricity. When two serially presented flash rates were compared, visual-visual matching judgments increased with eccentricity at flash rates of 6 Hz and higher, but decreased at flash rates below 6 Hz. Finally, when two concurrent flash rates were compared, visual-visual synchrony judgments increased with eccentricity at all flash-rate combinations. Together, these results suggest that while perceptual uncertainty can play a role in synchrony judgments at rates of 6 Hz and higher, below 6 Hz eccentricity produces a widening of the window of apparent audiovisual temporal synchrony which perceptual uncertainty cannot explain.

Acoustic Stimulation↗

Temporal relations of the complex spike activity of Purkinje cell pairs in the vestibulocerebellum of rabbits.

Parasagittal zones in the vestibulocerebellum contain Purkinje cells whose complex spike (CS) activity is modulated in response to rotational optokinetic stimulation (OKS) about either the vertical axis (VA) or a horizontal axis (HA) that is approximately perpendicular to the ipsilateral anterior canal. In rabbits, there are two VA zones in both the ventral nodulus and flocculus, two HA zones in the flocculus, and one HA zone in the ventral nodulus. We investigated the temporal relationship of the CS activity of Purkinje cell pairs in the same or different zones of the vestibulocerebellum in ketamine-anesthetized pigmented rabbits. A synchronous temporal relationship was defined as the tendency of the CS of each Purkinje cell to fire within, at most, 2 msec of one another. Generally, neurons in the same zone showed a tendency to exhibit CS synchrony. Of 82 pairs consisting of two Purkinje cells in the same zone (e.g., two nodulus HA cells), 33 were synchronous. In contrast, none of 26 pairs consisting of two neurons in functionally different zones (e.g., a VA cell paired with an HA cell), showed CS synchrony. Pairs consisting of neurons in spatially separated VA zones in the ventral nodulus also showed a tendency to be synchronously related (6/16), as did pairs consisting of a nodulus VA cell and a flocculus VA cell (3/14). The CS synchrony was higher during OKS in the preferred direction than during spontaneous activity. This is the first demonstration that CS synchrony in the vestibulocerebellum can be manipulated with a natural sensory stimulus.

Animals↗

Neural synergy in visual grouping: when good continuation meets common fate.

A modified version of the 'path finder' display consisting of many small oriented Gabor patches was used to study the joint contributions of spatial and temporal structures to shape perception. A two-interval forced-choice procedure measured detectability of curved 'paths' defined by orientation ('good continuation') and/or by temporal synchrony of change in motion direction ('common fate'). When orientation was completely random (no spatial 'path' cue) temporal synchrony still supported reliable performance, but only when correlation of change among 'path' elements was high. When combined, these two weak spatial and temporal structures yielded performance in excess of probability summation: 'paths' weakly defined by orientation were highly conspicuous when the constituent Gabors underwent synchronized changes in direction of motion, even though the individual directions of path elements were uncorrelated. Spatial grouping from temporal structure may arise from correlated transients associated with synchronized changes in motion direction. Evidently these two mechanisms for promotion of spatial grouping interact synergistically.

Binomial Distribution↗

Patterns of influenza-associated mortality among US elderly by geographic region and virus subtype, 1968-1998.

The regular seasonality of influenza in temperate countries is recognized, but regional differences in patterns of influenza-related mortality are poorly understood. Identifying patterns could improve epidemic prediction and prevention. The authors analyzed the monthly percentage of deaths attributable to pneumonia and influenza among people aged 65 or more years in the contiguous United States, 1968-1998. The local Moran's I test for spatial autocorrelation and correlograms assessing space-time synchrony within each influenza season were applied to detect and to characterize mortality patterns. Western US regions experienced epidemics of greater magnitude than did eastern regions. Positive spatial autocorrelation (two-sided p = 0.001) revealed the similarity in influenza mortality of neighboring states, with several western states forming a focus of high mortality. In transmission seasons dominated by virus subtype A(H3N2), mortality was correlated at a high and consistent level across the United States (mean correlation = 0.56, standard deviation = 0.134). However, when subtype A(H1N1) or type B dominated, the average synchrony was lower (mean correlation = 0.23, standard deviation = 0.058). These novel analyses suggest that causes of spatial heterogeneity (e.g., large-scale environmental drivers and population movement) have impacted influenza-associated mortality.

Aged↗

[Characteristics, dynamics and niche of insect community in plum orchard].

The insect community in plum orchard was investigated on organization level and temporal-spatial niche. The results showed that the insect community was abundant, which included 6 orders, 23 families. The individuals of species, diversity indices, and evenness increased with time. Myzus persicae and Asiaarposina sasokii had the widest spatial niche breadth, while Didesmococcus koreauus borchs had the widest temporal niche breadth. Among the natural enemies, Chilocorus rubidus had the widest both temporal and spatial niche breadth. The niche of Chicocorus rubidus and Didesmococcus koreauus overlapped larger than that of the others, which indicated their synchrony in temporal dimension and their similarity in spatial dimension. As the dominant natural enemies, the two populations should be protected and utilized to control plum pest.

Animals↗

Origin, structure, and role of background EEG activity. Part 1. Analytic amplitude.

OBJECTIVE: To explain the neural mechanisms of spontaneous EEG by measuring the spatiotemporal patterns of synchrony among beta-gamma oscillations during perception. METHODS: EEGs were measured from 8 x 8 (5.6 x 5.6 mm2) arrays fixed on the surfaces of primary sensory areas in rabbits that were trained to discriminate visual, auditory or tactile conditioned stimuli (CSs) eliciting conditioned responses (CRs). EEG preprocessing was by (i) bandpass filtering to extract the beta-gamma range (deleting theta-alpha); (ii) low-pass spatial filtering (not high-pass Laplacians used for localization), (iii) spatial averaging (not time averaging used for evoked potentials), and (iv) close spacing of 64 electrodes for simultaneous recording in each area (not sampling single signals from several areas); (v) novel algorithms were devised to measure synchrony and spatial pattern stability by calculating variances among patterns in 64-space derived from the 8 x 8 arrays (not by fitting equivalent dipoles). These methodological differences are crucial for the proposed new perspective on EEG. RESULTS: Spatial patterns of beta-gamma EEG emerged following sudden jumps in cortical activity called 'state transitions'. Each transition began with an abrupt phase re-setting to a new value on every channel, followed sequentially by re-synchronization, spatial pattern stabilization, and a dramatic increase in pattern amplitude. State transitions recurred at varying intervals in the theta range. A novel parameter was devised to estimate the perceptual information in the beta-gamma EEG, which disclosed 2-4 patterns with high information content in the CS-CR interval on each trial; each began with a state transition and lasted approximately 0.1 s. CONCLUSIONS: The function of each primary sensory neocortex was discontinuous; discrete spatial patterns occurred in frames like those in cinema. The frames before and after the CS-CR interval had low content. SIGNIFICANCE: Derivation and interpretation of unit data in studies of perception might benefit from using multichannel EEG recordings to define distinctive epochs that are demarcated by state transitions of neocortical dynamics in the CS-CR intervals, particularly in consideration of the possibility that EEG may reveal recurring episodes of exchange and sharing of perceptual information among multiple sensory cortices. Simultaneously recorded, multichannel beta-gamma EEG might assist in the interpretation of images derived by fMRI, since high beta-gamma EEG amplitudes imply high rates of energy utilization. The spatial pattern intermittency provides a tag to distinguish gamma bursts from contaminating EMG activity in scalp recording in order to establish beta-gamma recording as a standard clinical tool. Finally, EEG cannot fail to have a major impact on brain theory.

Acoustic Stimulation↗

Attention in natural scenes: Neurophysiological and computational bases.

How does attention operate in natural scenes? We show that the receptive fields of inferior temporal cortex neurons that implement object representations become small and located at the fovea in complex natural scenes. This facilitates the readout of information about an object that may be reward or punishment associated, and may be the target for action. Top-down biased competition to implement attention has a much weaker effect in complex natural scenes than in otherwise blank scenes with two objects. Part of the solution to the binding problem is thus that competition and the foveal cortical magnification factor emphasize what is present at the fovea, and limit the binding problem. Part of the solution to the binding problem is that neurons respond to combinations of features present in the correct relative spatial positions. Stimulus-dependent neuronal synchrony does not appear to be quantitatively important in feature binding, and in attention, in natural visual scenes, at least in the inferior temporal visual cortex, as shown by information theoretic analyses. The perception of multiple objects in a scene is facilitated by the fact that inferior temporal visual cortex neurons have asymmetrical receptive fields with respect to the fovea in complex scenes. Computational models of this processing are described.

Action Potentials↗

Gap junctions between interneuron dendrites can enhance synchrony of gamma oscillations in distributed networks.

Gamma-frequency (30-70 Hz) oscillations in populations of interneurons may be of functional relevance in the brain by virtue of their ability to induce synchronous firing in principal neurons. Such a role would require that neurons, 1 mm or more apart, be able to synchronize their activity, despite the presence of axonal conduction delays and of the limited axonal spread of many interneurons. We showed previously that interneuron doublet firing can help to synchronize gamma oscillations, provided that sufficiently many pyramidal neurons are active; we also suggested that gap junctions, between the axons of principal neurons, could contribute to the long-range synchrony of gamma oscillations induced in the hippocampus by carbachol in vitro. Here we consider interneuron network gamma: that is, gamma oscillations in pharmacologically isolated networks of tonically excited interneurons, with frequency gated by mutual GABA(A) receptor-mediated IPSPs. We provide simulation and electrophysiological evidence that interneuronal gap junctions (presumably dendritic) can enhance the synchrony of such gamma oscillations, in spatially extended interneuron networks. There appears to be a sharp threshold conductance, below which the interneuron dendritic gap junctions do not exert a synchronizing role.

Animals↗

Comprehension of Euclidean space in young children: the early emergence of understanding and its limits.

In two studies Ss (N = 114) aged 3-5 years of age were assessed as to their ability to reconstruct or recognize spatial displays depicting Euclidean geometric features. The Ss responded either with the standard display present or in its absence, a memory condition. Items differed as to whether the objects located in the plane required attention to either one axis, the horizontal or vertical, or to both. In Study I Ss constructed their matches with an open-ended assessment device, while in Study II, they selected the match from a field of four distractors. Results from both studies were convergent despite differences in assessment technique and specifics of data analysis, and indicated an effect for age and condition, and that children do less well when dealing with double-axis displays in the memory condition. In addition, it appears that males were less attentive to display orientation cues than were females. Results are discussed in terms of Piagetian theoretical formulations about children's spatial development, their sensitivity to Euclidean spatial features, and the lack of synchrony between perceptual abilities and memory.

Attention↗

Gamma oscillation maintains stimulus structure-dependent synchronization in cat visual cortex.

Visual cortical cells demonstrate both oscillation and synchronization, although the underlying causes and functional significance of these behaviors remain uncertain. We simultaneously recorded single-unit activity with microelectrode arrays in supragranular layers of area 17 of cats paralyzed and anesthetized with propofol and N(2)O. Rate-normalized autocorrelograms of 24 cells reveal bursting (100%) and gamma oscillation (63%). Renewal density analysis, used to explore the source of oscillation, suggests a contribution from extrinsic influences such as feedback. However, a bursting refractory period, presumably membrane-based, could also encourage oscillatory firing. When we investigated the source of synchronization for 60 cell pairs we found only moderate correlation of synchrony with bursts and oscillation. We did, nonetheless, discover a possible functional role for oscillation. In all cases of cross-correlograms that exhibited oscillation, the strength of the synchrony was maintained throughout the stimulation period. When no oscillation was apparent, 75% of the cell pairs showed decay in their synchronization. The synchrony between cells is strongly dependent on similar response onset latencies. We therefore propose that structured input, which yields tight organization of latency, is a more likely candidate for the source of synchronization than oscillation. The reliable synchrony at response onset could be driven by spatial and temporal correlation of the stimulus that is preserved through the earlier stages of the visual system. Oscillation then contributes to maintenance of the synchrony to enhance reliable transmission of the information for higher cognitive processing.

Action Potentials↗