A neural mass model of place cell activity: theta phase precession, replay and imagination of never experienced paths.

Recent results on hippocampal place cells show that the replay of behavioral sequences does not simply reflect previously experienced trajectories, but may also occur in the reverse direction, or may even include never experienced paths. In order to elucidate the possible mechanisms at the basis of this phenomenon, we have developed a model of sequence learning. The present model consists of two layers of place cell units. Long-range connections among units implement heteroassociation between the two layers, trained with a temporal Hebb rule. The network was trained assuming that a virtual rat moves within a virtual maze. This training leads to the formation of bidirectional synapses between the two layers, i.e. synapses connecting a neuron both with its previous and subsequent element in the path. Subsequently, two distinct conditions were simulated with the trained network. During an exploratory phase, characterized by a similar consideration to the external environment and to the internal representation, the model simulates the occurrence of theta precession in the forward path and the temporal compression. During an imagination phase, when there is no consideration to the external location, the model produces trains of gamma oscillations, without the presence of a theta rhythm, and simulates the occurrence of both direct and reverse replay, and the imagination of never experienced paths. The new paths are built by combining bunches of previous trajectories. The main mechanisms at the basis of this behavior are explained in detail, and lines for future improvements (e.g., to simulate preplay) are discussed.

An improved algorithm for the automatic detection and characterization of slow eye movements.

Slow eye movements (SEMs) are typical of drowsy wakefulness and light sleep. SEMs still lack of systematic physical characterization. We present a new algorithm, which substantially improves our previous one, for the automatic detection of SEMs from the electro-oculogram (EOG) and extraction of SEMs physical parameters. The algorithm utilizes discrete wavelet decomposition of the EOG to implement a Bayes classifier that identifies intervals of slow ocular activity; each slow activity interval is segmented into single SEMs via a template matching method. Parameters of amplitude, duration, velocity are automatically extracted from each detected SEM. The algorithm was trained and validated on sleep onsets and offsets of 20 EOG recordings visually inspected by an expert. Performances were assessed in terms of correctly identified slow activity epochs (sensitivity: 85.12%; specificity: 82.81%), correctly segmented single SEMs (89.08%), and time misalignment (0.49 s) between the automatically and visually identified SEMs. The algorithm proved reliable even in whole sleep (sensitivity: 83.40%; specificity: 72.08% in identifying slow activity epochs; correctly segmented SEMs: 93.24%; time misalignment: 0.49 s). The algorithm, being able to objectively characterize single SEMs, may be a valuable tool to improve knowledge of normal and pathological sleep.

A neural network model can explain ventriloquism aftereffect and its generalization across sound frequencies.

Exposure to synchronous but spatially disparate auditory and visual stimuli produces a perceptual shift of sound location towards the visual stimulus (ventriloquism effect). After adaptation to a ventriloquism situation, enduring sound shift is observed in the absence of the visual stimulus (ventriloquism aftereffect). Experimental studies report opposing results as to aftereffect generalization across sound frequencies varying from aftereffect being confined to the frequency used during adaptation to aftereffect generalizing across some octaves. Here, we present an extension of a model of visual-auditory interaction we previously developed. The new model is able to simulate the ventriloquism effect and, via Hebbian learning rules, the ventriloquism aftereffect and can be used to investigate aftereffect generalization across frequencies. The model includes auditory neurons coding both for the spatial and spectral features of the auditory stimuli and mimicking properties of biological auditory neurons. The model suggests that different extent of aftereffect generalization across frequencies can be obtained by changing the intensity of the auditory stimulus that induces different amounts of activation in the auditory layer. The model provides a coherent theoretical framework to explain the apparently contradictory results found in the literature. Model mechanisms and hypotheses are discussed in relation to neurophysiological and psychophysical data.

A multi-layer neural-mass model for learning sequences using theta/gamma oscillations.

A neural mass model for the memorization of sequences is presented. It exploits three layers of cortical columns that generate a theta/gamma rhythm. The first layer implements an auto-associative memory working in the theta range; the second segments objects in the gamma range; finally, the feedback interactions between the third and the second layers realize a hetero-associative memory for learning a sequence. After training with Hebbian and anti-Hebbian rules, the network recovers sequences and accounts for the phase-precession phenomenon.

Binding and segmentation via a neural mass model trained with Hebbian and anti-Hebbian mechanisms.

Synchronization of neural activity in the gamma band, modulated by a slower theta rhythm, is assumed to play a significant role in binding and segmentation of multiple objects. In the present work, a recent neural mass model of a single cortical column is used to analyze the synaptic mechanisms which can warrant synchronization and desynchronization of cortical columns, during an autoassociation memory task. The model considers two distinct layers communicating via feedforward connections. The first layer receives the external input and works as an autoassociative network in the theta band, to recover a previously memorized object from incomplete information. The second realizes segmentation of different objects in the gamma band. To this end, units within both layers are connected with synapses trained on the basis of previous experience to store objects. The main model assumptions are: (i) recovery of incomplete objects is realized by excitatory synapses from pyramidal to pyramidal neurons in the same object; (ii) binding in the gamma range is realized by excitatory synapses from pyramidal neurons to fast inhibitory interneurons in the same object. These synapses (both at points i and ii) have a few ms dynamics and are trained with a Hebbian mechanism. (iii) Segmentation is realized with faster AMPA synapses, with rise times smaller than 1 ms, trained with an anti-Hebbian mechanism. Results show that the model, with the previous assumptions, can correctly reconstruct and segment three simultaneous objects, starting from incomplete knowledge. Segmentation of more objects is possible but requires an increased ratio between the theta and gamma periods.

The generation of rhythms within a cortical region: analysis of a neural mass model.

Rhythms in brain electrical activity are assumed to play a significant role in many cognitive and perceptual processes. It is thus of great value to analyze these rhythms and their mutual relationships in large scale models of cortical regions. In the present work, we modified the neural mass model by Wendling et al. (Eur. J. Neurosci. 15 (2002) 1499-1508) by including a new inhibitory self-loop among GABAA,fast interneurons. A theoretical analysis was performed to demonstrate that, thanks to this loop, GABAA,fast interneurons can produce a gamma rhythm in the power spectral density (PSD) even without the participation of the other neural populations. Then, the model of a whole cortical region, built upon four interconnected neural populations (pyramidal cells, excitatory, GABAA,slow and GABAA,fast interneurons) was investigated by changing the internal connectivity parameters. Results show that different rhythm combinations (beta and gamma, alpha and gamma, or a wide spectrum) can be obtained within the same region by simply altering connectivity values, without the need to change synaptic kinetics. Finally, two or three cortical regions were connected by using different topologies of long range connections. Results show that long-range connections directed from pyramidal neurons to GABAA,fast interneurons are the most efficient to transmit rhythms from one region to another. In this way, PSD with three or four peaks can be obtained using simple connectivity patterns. The model can be of value to gain a deeper insight into the mechanisms involved in the generation of gamma rhythms and provide a better understanding of cortical EEG spectra.

Changes in EEG power spectral density and cortical connectivity in healthy and tetraplegic patients during a motor imagery task.

Knowledge of brain connectivity is an important aspect of modern neuroscience, to understand how the brain realizes its functions. In this work, neural mass models including four groups of excitatory and inhibitory neurons are used to estimate the connectivity among three cortical regions of interests (ROIs) during a foot-movement task. Real data were obtained via high-resolution scalp EEGs on two populations: healthy volunteers and tetraplegic patients. A 3-shell Boundary Element Model of the head was used to estimate the cortical current density and to derive cortical EEGs in the three ROIs. The model assumes that each ROI can generate an intrinsic rhythm in the beta range, and receives rhythms in the alpha and gamma ranges from other two regions. Connectivity strengths among the ROIs were estimated by means of an original genetic algorithm that tries to minimize several cost functions of the difference between real and model power spectral densities. Results show that the stronger connections are those from the cingulate cortex to the primary and supplementary motor areas, thus emphasizing the pivotal role played by the CMA_L during the task. Tetraplegic patients exhibit higher connectivity strength on average, with significant statistical differences in some connections. The results are commented and virtues and limitations of the proposed method discussed.