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A small world of neuronal synchrony
(2008)
- A small-world network has been suggested to be an efficient solution for achieving both modular and global processing-a property highly desirable for brain computations. Here, we investigated functional networks of cortical neurons using correlation analysis to identify functional connectivity. To reconstruct the interaction network, we applied the Ising model based on the principle of maximum entropy. This allowed us to assess the interactions by measuring pairwise correlations and to assess the strength of coupling from the degree of synchrony. Visual responses were recorded in visual cortex of anesthetized cats, simultaneously from up to 24 neurons. First, pairwise correlations captured most of the patterns in the population´s activity and, therefore, provided a reliable basis for the reconstruction of the interaction networks. Second, and most importantly, the resulting networks had small-world properties; the average path lengths were as short as in simulated random networks, but the clustering coefficients were larger. Neurons differed considerably with respect to the number and strength of interactions, suggesting the existence of "hubs" in the network. Notably, there was no evidence for scale-free properties. These results suggest that cortical networks are optimized for the coexistence of local and global computations: feature detection and feature integration or binding.
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Analyzing possible pitfalls of cross-frequency analysis : poster presentation from Twentieth Annual Computational Neuroscience Meeting CNS*2011 Stockholm, Sweden, 23 - 28 July 2011
(2011)
- Poster presentation from Twentieth Annual Computational Neuroscience Meeting: CNS*2011 Stockholm, Sweden. 23-28 July 2011. One of the central questions in neuroscience is how neural activity is organized across different spatial and temporal scales. As larger populations oscillate and synchronize at lower frequencies and smaller ensembles are active at higher frequencies, a cross-frequency coupling would facilitate flexible coordination of neural activity simultaneously in time and space. Although various experiments have revealed amplitude-to-amplitude and phase-to-phase coupling, the most common and most celebrated result is that the phase of the lower frequency component modulates the amplitude of the higher frequency component. Over the recent 5 years the amount of experimental works finding such phase-amplitude coupling in LFP, ECoG, EEG and MEG has been tremendous (summarized in [1]). We suggest that although the mechanism of cross-frequency-coupling (CFC) is theoretically very tempting, the current analysis methods might overestimate any physiological CFC actually evident in the signals of LFP, ECoG, EEG and MEG. In particular, we point out three conceptual problems in assessing the components and their correlations of a time series. Although we focus on phase-amplitude coupling, most of our argument is relevant for any type of coupling. 1) The first conceptual problem is related to isolating physiological frequency components of the recorded signal. The key point is to notice that there are many different mathematical representations for a time series but the physical interpretation we make out of them is dependent on the choice of the components to be analyzed. In particular, when one isolates the components by Fourier-representation based filtering, it is the width of the filtering bands what defines what we consider as our components and how their power or group phase change in time. We will discuss clear cut examples where the interpretation of the existence of CFC depends on the width of the filtering process. 2) A second problem deals with the origin of spectral correlations as detected by current cross-frequency analysis. It is known that non-stationarities are associated with spectral correlations in the Fourier space. Therefore, there are two possibilities regarding the interpretation of any observed CFC. One scenario is that basic neuronal mechanisms indeed generate an interaction across different time scales (or frequencies) resulting in processes with non-stationary features. The other and problematic possibility is that unspecific non-stationarities can also be associated with spectral correlations which in turn will be detected by cross frequency measures even if physiologically there is no causal interaction between the frequencies. 3) We discuss on the role of non-linearities as generators of cross frequency interactions. As an example we performed a phase-amplitude coupling analysis of two nonlinearly related signals: atmospheric noise and the square of it (Figure 1) observing an enhancement of phase-amplitude coupling in the second signal while no pattern is observed in the first. Finally, we discuss some minimal conditions need to be tested to solve some of the ambiguities here noted. In summary, we simply want to point out that finding a significant cross frequency pattern does not always have to imply that there indeed is physiological cross frequency interaction in the brain.
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Auditory motion capturing ambiguous visual motion
(2012)
- In this study, it is demonstrated that moving sounds have an effect on the direction in which one sees visual stimuli move. During the main experiment sounds were presented consecutively at four speaker locations inducing left or rightward auditory apparent motion. On the path of auditory apparent motion, visual apparent motion stimuli were presented with a high degree of directional ambiguity. The main outcome of this experiment is that our participants perceived visual apparent motion stimuli that were ambiguous (equally likely to be perceived as moving left or rightward) more often as moving in the same direction than in the opposite direction of auditory apparent motion. During the control experiment we replicated this finding and found no effect of sound motion direction on eye movements. This indicates that auditory motion can capture our visual motion percept when visual motion direction is insufficiently determinate without affecting eye movements.
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Callosal connections of primary visual cortex predict the spatial spreading of binocular rivalry across the visual hemifields
(2011)
- In binocular rivalry, presentation of different images to the separate eyes leads to conscious perception alternating between the two possible interpretations every few seconds. During perceptual transitions, a stimulus emerging into dominance can spread in a wave-like manner across the visual field. These traveling waves of rivalry dominance have been successfully related to the cortical magnification properties and functional activity of early visual areas, including the primary visual cortex (V1). Curiously however, these traveling waves undergo a delay when passing from one hemifield to another. In the current study, we used diffusion tensor imaging (DTI) to investigate whether the strength of interhemispheric connections between the left and right visual cortex might be related to the delay of traveling waves across hemifields. We measured the delay in traveling wave times (ΔTWT) in 19 participants and repeated this test 6 weeks later to evaluate the reliability of our behavioral measures. We found large interindividual variability but also good test–retest reliability for individual measures of ΔTWT. Using DTI in connection with fiber tractography, we identified parts of the corpus callosum connecting functionally defined visual areas V1–V3. We found that individual differences in ΔTWT was reliably predicted by the diffusion properties of transcallosal fibers connecting left and right V1, but observed no such effect for neighboring transcallosal visual fibers connecting V2 and V3. Our results demonstrate that the anatomical characteristics of topographically specific transcallosal connections predict the individual delay of interhemispheric traveling waves, providing further evidence that V1 is an important site for neural processes underlying binocular rivalry.
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Context Matters: The Illusive Simplicity of Macaque V1 Receptive Fields
(2012)
- Even in V1, where neurons have well characterized classical receptive fields (CRFs), it has been difficult to deduce which features of natural scenes stimuli they actually respond to. Forward models based upon CRF stimuli have had limited success in predicting the response of V1 neurons to natural scenes. As natural scenes exhibit complex spatial and temporal correlations, this could be due to surround effects that modulate the sensitivity of the CRF. Here, instead of attempting a forward model, we quantify the importance of the natural scenes surround for awake macaque monkeys by modeling it non-parametrically. We also quantify the influence of two forms of trial to trial variability. The first is related to the neuron’s own spike history. The second is related to ongoing mean field population activity reflected by the local field potential (LFP). We find that the surround produces strong temporal modulations in the firing rate that can be both suppressive and facilitative. Further, the LFP is found to induce a precise timing in spikes, which tend to be temporally localized on sharp LFP transients in the gamma frequency range. Using the pseudo R2 as a measure of model fit, we find that during natural scene viewing the CRF dominates, accounting for 60% of the fit, but that taken collectively the surround, spike history and LFP are almost as important, accounting for 40%. However, overall only a small proportion of V1 spiking statistics could be explained (R2~5%), even when the full stimulus, spike history and LFP were taken into account. This suggests that under natural scene conditions, the dominant influence on V1 neurons is not the stimulus, nor the mean field dynamics of the LFP, but the complex, incoherent dynamics of the network in which neurons are embedded.
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Deficits in high- (>60 Hz) gamma-band oscillations during visual processing in schizophrenia
(2013)
- Current theories of the pathophysiology of schizophrenia have focused on abnormal temporal coordination of neural activity. Oscillations in the gamma-band range (>25 Hz) are of particular interest as they establish synchronization with great precision in local cortical networks. However, the contribution of high gamma (>60 Hz) oscillations toward the pathophysiology is less established. To address this issue, we recorded magnetoencephalographic (MEG) data from 16 medicated patients with chronic schizophrenia and 16 controls during the perception of Mooney faces. MEG data were analysed in the 25–150 Hz frequency range. Patients showed elevated reaction times and reduced detection rates during the perception of upright Mooney faces while responses to inverted stimuli were intact. Impaired processing of Mooney faces in schizophrenia patients was accompanied by a pronounced reduction in spectral power between 60–120 Hz (effect size: d = 1.26) which was correlated with disorganized symptoms (r = −0.72). Our findings demonstrate that deficits in high gamma-band oscillations as measured by MEG are a sensitive marker for aberrant cortical functioning in schizophrenia, suggesting an important aspect of the pathophysiology of the disorder.
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Detecting multineuronal temporal patterns in parallel spike trains
(2012)
- We present a non-parametric and computationally efficient method that detects spatiotemporal firing patterns and pattern sequences in parallel spike trains and tests whether the observed numbers of repeating patterns and sequences on a given timescale are significantly different from those expected by chance. The method is generally applicable and uncovers coordinated activity with arbitrary precision by comparing it to appropriate surrogate data. The analysis of coherent patterns of spatially and temporally distributed spiking activity on various timescales enables the immediate tracking of diverse qualities of coordinated firing related to neuronal state changes and information processing. We apply the method to simulated data and multineuronal recordings from rat visual cortex and show that it reliably discriminates between data sets with random pattern occurrences and with additional exactly repeating spatiotemporal patterns and pattern sequences. Multineuronal cortical spiking activity appears to be precisely coordinated and exhibits a sequential organization beyond the cell assembly concept.
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Die Welt jenseits der Oszillografen : ein Streitgespräch zwischen dem Hirnforscher Wolf Singer und dem Philosophen Marcus Willaschek
(2005)
- Neurowissenschaftler fordern einen illusionslosen Umgang mit Begriffen wie Willensfreiheit und Bewusstsein. Philosophen kritisieren offen die Thesen von Hirnforschern. Stehen sich diese Positionen unversöhnlich gegenüber? Wo gibt es Möglichkeiten einer Annäherung, gar einer Kooperation? Der Religionsphilosoph Prof. Dr. Thomas M. Schmidt und der Biologe Stefan Kieß loten die Situation in Frankfurt aus; ihre Gesprächspartner sind der Hirnforscher Prof. Dr. Wolf Singer (links), Direktor am Max-Planck-Institut für Hirnforschung, und Prof. Dr. Marcus Willaschek (rechts), Philosoph an der Universität Frankfurt.
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Distributed fading memory for stimulus properties in the primary visual cortex
(2009)
- It is currently not known how distributed neuronal responses in early visual areas carry stimulus-related information. We made multielectrode recordings from cat primary visual cortex and applied methods from machine learning in order to analyze the temporal evolution of stimulus-related information in the spiking activity of large ensembles of around 100 neurons. We used sequences of up to three different visual stimuli (letters of the alphabet) presented for 100 ms and with intervals of 100 ms or larger. Most of the information about visual stimuli extractable by sophisticated methods of machine learning, i.e., support vector machines with nonlinear kernel functions, was also extractable by simple linear classification such as can be achieved by individual neurons. New stimuli did not erase information about previous stimuli. The responses to the most recent stimulus contained about equal amounts of information about both this and the preceding stimulus. This information was encoded both in the discharge rates (response amplitudes) of the ensemble of neurons and, when using short time constants for integration (e.g., 20 ms), in the precise timing of individual spikes (<= ~20 ms), and persisted for several 100 ms beyond the offset of stimuli. The results indicate that the network from which we recorded is endowed with fading memory and is capable of performing online computations utilizing information about temporally sequential stimuli. This result challenges models assuming frame-by-frame analyses of sequential inputs.
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Distributed processing and temporal codes in neuronal networks
(2009)
- The cerebral cortex presents itself as a distributed dynamical system with the characteristics of a small world network. The neuronal correlates of cognitive and executive processes often appear to consist of the coordinated activity of large assemblies of widely distributed neurons. These features require mechanisms for the selective routing of signals across densely interconnected networks, the flexible and context dependent binding of neuronal groups into functionally coherent assemblies and the task and attention dependent integration of subsystems. In order to implement these mechanisms, it is proposed that neuronal responses should convey two orthogonal messages in parallel. They should indicate (1) the presence of the feature to which they are tuned and (2) with which other neurons (specific target cells or members of a coherent assembly) they are communicating. The first message is encoded in the discharge frequency of the neurons (rate code) and it is proposed that the second message is contained in the precise timing relationships between individual spikes of distributed neurons (temporal code). It is further proposed that these precise timing relations are established either by the timing of external events (stimulus locking) or by internal timing mechanisms. The latter are assumed to consist of an oscillatory modulation of neuronal responses in different frequency bands that cover a broad frequency range from <2 Hz (delta) to >40 Hz (gamma) and ripples. These oscillations limit the communication of cells to short temporal windows whereby the duration of these windows decreases with oscillation frequency. Thus, by varying the phase relationship between oscillating groups, networks of functionally cooperating neurons can be flexibly configurated within hard wired networks. Moreover, by synchronizing the spikes emitted by neuronal populations, the saliency of their responses can be enhanced due to the coincidence sensitivity of receiving neurons in very much the same way as can be achieved by increasing the discharge rate. Experimental evidence will be reviewed in support of the coexistence of rate and temporal codes. Evidence will also be provided that disturbances of temporal coding mechanisms are likely to be one of the pathophysiological mechanisms in schizophrenia.
