Memory and forgetting constitute two sides of the same coin, and although the first has been extensively investigated, the latter is often overlooked. A possible approach to better understand forgetting is to develop phenomenological models that implement its putative mechanisms in the most elementary way possible, and then experimentally test the theoretical predictions of these models. One such mechanism proposed in previous studies is retrograde interference, stating that a memory can be erased due to subsequently acquired memories. In the current contribution, we hypothesize that retrograde erasure is controlled by the relevant “importance” measures such that more important memories eliminate less important ones acquired earlier. We show that some versions of the resulting mathematical model are broadly compatible with the previously reported power-law forgetting time course and match well the results of our recognition experiments with long, randomly assembled streams of words.
Decoding approaches provide a useful means of estimating the information contained in neuronal circuits. In this work, we analyze the expected classification error of a decoder based on Fisher linear discriminant analysis. We provide expressions that relate decoding error to the specific parameters of a population model that performs linear integration of sensory input. Results show conditions that lead to beneficial and detrimental effects of noise correlation on decoding. Further, the proposed framework sheds light on the contribution of neuronal noise, highlighting cases where, counter-intuitively, increased noise may lead to improved decoding performance. Finally, we examined the impact of dynamical parameters, including neuronal leak and integration time constant, on decoding. Overall, this work presents a fruitful approach to the study of decoding using a comprehensive theoretical framework that merges dynamical parameters with estimates of readout error.
The brain is intrinsically organized into large-scale networks that constantly re-organize on multiple timescales, even when the brain is at rest. The timing of these dynamics is crucial for sensation, perception, cognition, and ultimately consciousness, but the underlying dynamics governing the constant reorganization and switching between networks are not yet well understood. Electroencephalogram (EEG) microstates are brief periods of stable scalp topography that have been identified as the electrophysiological correlate of functional magnetic resonance imaging defined resting-state networks. Spatiotemporal microstate sequences maintain high temporal resolution and have been shown to be scale-free with long-range temporal correlations. Previous attempts to model EEG microstate sequences have failed to capture this crucial property and so cannot fully capture the dynamics; this paper answers the call for more sophisticated modeling approaches. We present a dynamical model that exhibits a noisy network attractor between nodes that represent the microstates. Using an excitable network between four nodes, we can reproduce the transition probabilities between microstates but not the heavy tailed residence time distributions. We present two extensions to this model: first, an additional hidden node at each state; second, an additional layer that controls the switching frequency in the original network. Introducing either extension to the network gives the flexibility to capture these heavy tails. We compare the model generated sequences to microstate sequences from EEG data collected from healthy subjects at rest. For the first extension, we show that the hidden nodes 'trap' the trajectories allowing the control of residence times at each node. For the second extension, we show that two nodes in the controlling layer are sufficient to model the long residence times. Finally, we show that in addition to capturing the residence time distributions and transition probabilities of the sequences, these two models capture additional properties of the sequences including having interspersed long and short residence times and long range temporal correlations in line with the data as measured by the Hurst exponent.
The principle of stationary action is a cornerstone of modern physics, providing a powerful framework for investigating dynamical systems found in classical mechanics through to quantum field theory. However, computational neuroscience, despite its heavy reliance on concepts in physics, is anomalous in this regard as its main equations of motion are not compatible with a Lagrangian formulation and hence with the principle of stationary action. Taking the Dynamic Causal Modelling (DCM) neuronal state equation as an instructive archetype of the first-order linear differential equations commonly found in computational neuroscience, we show that it is possible to make certain modifications to this equation to render it compatible with the principle of stationary action. Specifically, we show that a Lagrangian formulation of the DCM neuronal state equation is facilitated using a complex dependent variable, an oscillatory solution, and a Hermitian intrinsic connectivity matrix. We first demonstrate proof of principle by using Bayesian model inversion to show that both the original and modified models can be correctly identified via in silico data generated directly from their respective equations of motion. We then provide motivation for adopting the modified models in neuroscience by using three different types of publicly available in vivo neuroimaging datasets, together with open source MATLAB code, to show that the modified (oscillatory) model provides a more parsimonious explanation for some of these empirical timeseries. It is our hope that this work will, in combination with existing techniques, allow people to explore the symmetries and associated conservation laws within neural systems – and to exploit the computational expediency facilitated by direct variational techniques.
We report two cases of eccrine syringofibroadenoma. Both patients had a large, solitary hyperkeratotic nodular lesion over the extremities. Histologic sections showed spongelike masses of small cuboidal acrosyringeal cells. Fibrovascular connective tissue stroma resembling premalignant fibroepithelial tumor of Pinkus filled the spaces between the masses.
Mathematical models at multiple temporal and spatial scales can unveil the fundamental mechanisms of critical transitions in brain activities. Neural mass models (NMMs) consider the average temporal dynamics of interconnected neuronal subpopulations without explicitly representing the underlying cellular activity. The mesoscopic level offered by the neural mass formulation has been used to model electroencephalographic (EEG) recordings and to investigate various cerebral mechanisms, such as the generation of physiological and pathological brain activities. In this work, we consider a NMM widely accepted in the context of epilepsy, which includes four interacting neuronal subpopulations with different synaptic kinetics. Due to the resulting three-time-scale structure, the model yields complex oscillations of relaxation and bursting types. By applying the principles of geometric singular perturbation theory, we unveil the existence of the canard solutions and detail how they organize the complex oscillations and excitability properties of the model. In particular, we show that boundaries between pathological epileptic discharges and physiological background activity are determined by the canard solutions. Finally we report the existence of canard-mediated small-amplitude frequency-specific oscillations in simulated local field potentials for decreased inhibition conditions. Interestingly, such oscillations are actually observed in intracerebral EEG signals recorded in epileptic patients during pre-ictal periods, close to seizure onsets.
We study pattern formation in a 2-population homogenized neural field model of the Hopfield type in one spatial dimension with periodic microstructure. The connectivity functions are periodically modulated in both the synaptic footprint and in the spatial scale. It is shown that the nonlocal synaptic interactions promote a finite band width instability. The stability method relies on a sequence of wave-number dependent invariants of $2\times 2$ -stability matrices representing the sequence of Fourier-transformed linearized evolution equations for the perturbation imposed on the homogeneous background. The generic picture of the instability structure consists of a finite set of well-separated gain bands. In the shallow firing rate regime the nonlinear development of the instability is determined by means of the translational invariant model with connectivity kernels replaced with the corresponding period averaged connectivity functions. In the steep firing rate regime the pattern formation process depends sensitively on the spatial localization of the connectivity kernels: For strongly localized kernels this process is determined by the translational invariant model with period averaged connectivity kernels, whereas in the complementary regime of weak and moderate localization requires the homogenized model as a starting point for the analysis. We follow the development of the instability numerically into the nonlinear regime for both steep and shallow firing rate functions when the connectivity kernels are modeled by means of an exponentially decaying function. We also study the pattern forming process numerically as a function of the heterogeneity parameters in four different regimes ranging from the weakly modulated case to the strongly heterogeneous case. For the weakly modulated regime, we observe that stable spatial oscillations are formed in the steep firing rate regime, whereas we get spatiotemporal oscillations in the shallow regime of the firing rate functions.
We analyse the potential effects of lateral connectivity (amacrine cells and gap junctions) on motion anticipation in the retina. Our main result is that lateral connectivity can—under conditions analysed in the paper—trigger a wave of activity enhancing the anticipation mechanism provided by local gain control (Berry et al. in Nature 398(6725):334–338, 1999; Chen et al. in J. Neurosci. 33(1):120–132, 2013). We illustrate these predictions by two examples studied in the experimental literature: differential motion sensitive cells (Baccus and Meister in Neuron 36(5):909–919, 2002) and direction sensitive cells where direction sensitivity is inherited from asymmetry in gap junctions connectivity (Trenholm et al. in Nat. Neurosci. 16:154–156, 2013). We finally present reconstructions of retinal responses to 2D visual inputs to assess the ability of our model to anticipate motion in the case of three different 2D stimuli.
In this work, we consider a general conductance-based neuron model with the inclusion of the acetycholine sensitive, M-current. We study bifurcations in the parameter space consisting of the applied current, $I_{app}$ the maximal conductance of the M-current, $g_M$, and the conductance of the leak current, $g_L$. We give precise conditions for the model that ensure the existence of a Bogdanov-Takens (BT) point and show such a point can occur by varying $I_{app}$ and $g_{M}$. We discuss the case when the BT point becomes a Bogdanov-Takens-Cusp (BTC) point and show that such a point can occur in the three dimensional parameter space. The results of the bifurcation analysis are applied to different neuronal models and are verified and supplemented by numerical bifurcation diagrams generated using the package MATCONT. We conclude that there is a transition in the neuronal excitability type organized by the BT point and the neuron switches from Class-I to Class-II as conductance of the M-current increases.
In the auditory streaming paradigm, alternating sequences of pure tones can be perceived as a single galloping rhythm (integration) or as two sequences with separated low and high tones (segregation). Although studied for decades, the neural mechanisms underlining this perceptual grouping of sound remains a mystery. With the aim of identifying a plausible minimal neural circuit that captures this phenomenon, we propose a firing rate model with two periodically forced neural populations coupled by fast direct excitation and slow delayed inhibition. By analyzing the model in a non-smooth, slow-fast regime we analytically prove the existence of a rich repertoire of dynamical states and of their parameter dependent transitions. We impose plausible parameter restrictions and link all states with perceptual interpretations. Regions of stimulus parameters occupied by states linked with each percept match those found in behavioural experiments. Our model suggests that slow inhibition masks the perception of subsequent tones during segregation (forward masking), whereas fast excitation enables integration for large pitch differences between the two tones.
The reconstruction mechanisms built by the human auditory system during sound reconstruction are still a matter of debate. The purpose of this study is to propose a mathematical model of sound reconstruction based on the functional architecture of the auditory cortex (A1). The model is inspired by the geometrical modelling of vision, which has undergone a great development in the last ten years. There are, however, fundamental dissimilarities, due to the different role played by time and the different group of symmetries. The algorithm transforms the degraded sound in an ‘image’ in the time–frequency domain via a short-time Fourier transform. Such an image is then lifted to the Heisenberg group and is reconstructed via a Wilson–Cowan integro-differential equation. Preliminary numerical experiments are provided, showing the good reconstruction properties of the algorithm on synthetic sounds concentrated around two frequencies.
AbstractWhite matter pathways form a complex network of myelinated axons that regulate signal transmission in the nervous system and play a key role in behaviour and cognition. Recent evidence reveals that white matter networks are adaptive and that myelin remodels itself in an activity-dependent way, during both developmental stages and later on through behaviour and learning. As a result, axonal conduction delays continuously adjust in order to regulate the timing of neural signals propagating between different brain areas. This delay plasticity mechanism has yet to be integrated in computational neural models, where conduction delays are oftentimes constant or simply ignored. As a first approach to adaptive white matter remodelling, we modified the canonical Kuramoto model by enabling all connections with adaptive, phase-dependent delays. We analyzed the equilibria and stability of this system, and applied our results to two oscillator and large dimensional networks. Our joint mathematical and numerical analysis demonstrates that plastic delays act as a stabilizing mechanism promoting the network’s ability to maintain synchronous activity. Our work also shows that global synchronization is more resilient to perturbations and injury towards network architecture. Our results provide key insights about the analysis and potential significance of activity-dependent myelination in large-scale brain synchrony.
The modeling of neural fields in the visual cortex involves geometrical structures which describe in mathematical formalism the functional architecture of this cortical area. The case of contour detection and orientation tuning has been extensively studied and has become a paradigm for the mathematical analysis of image processing by the brain. Ten years ago an attempt was made to extend these models by replacing orientation (an angle) with a second-order tensor built from the gradient of the image intensity, and it was named the structure tensor. This assumption does not follow from biological observations (experimental evidence is still lacking) but from the idea that the effectiveness of texture processing with the structure tensor in computer vision may well be exploited by the brain itself. The drawback is that in this case the geometry is not Euclidean but hyperbolic instead, which complicates the analysis substantially. The purpose of this review is to present the methodology that was developed in a series of papers to investigate this quite unusual problem, specifically from the point of view of tuning and pattern formation. These methods, which rely on bifurcation theory with symmetry in the hyperbolic context, might be of interest for the modeling of other features such as color vision or other brain functions.
Coarse-graining microscopic models of biological neural networks to obtain mesoscopic models of neural activities is an essential step towards multi-scale models of the brain. Here, we extend a recent theory for mesoscopic population dynamics with static synapses to the case of dynamic synapses exhibiting short-term plasticity (STP). The extended theory offers an approximate mean-field dynamics for the synaptic input currents arising from populations of spiking neurons and synapses undergoing Tsodyks-Markram STP. The approximate mean-field dynamics accounts for both finite number of synapses and correlation between the two synaptic variables of the model (utilization and available resources) and its numerical implementation is simple. Comparisons with Monte Carlo simulations of the microscopic model show that in both feedforward and recurrent networks, the mesoscopic mean-field model accurately reproduces the first- and second-order statistics of the total synaptic input into a postsynaptic neuron and accounts for stochastic switches between Up and Down states and for population spikes. The extended mesoscopic population theory of spiking neural networks with STP may be useful for a systematic reduction of detailed biophysical models of cortical microcircuits to numerically efficient and mathematically tractable mean-field models.
Essential tremor manifests predominantly as a tremor of the upper limbs. One therapy option is high-frequency deep brain stimulation, which continuously delivers electrical stimulation to the ventral intermediate nucleus of the thalamus at about 130 Hz. Investigators have been looking at stimulating less, chiefly to reduce side effects. One strategy, phase-locked deep brain stimulation, consists of stimulating according to the phase of the tremor, once per period. In this study, we aim to reproduce the phase dependent effects of stimulation seen in patient data with a biologically inspired Wilson-Cowan model. To this end, we first analyse patient data, and conclude that half of the datasets have response curves that are better described by sinusoidal curves than by straight lines, while an effect of phase cannot be consistently identified in the remaining half. Using the Hilbert phase we derive analytical expressions for phase and amplitude responses to phase-dependent stimulation and study their relationship in the linearisation of a stable focus model, a simplification of the Wilson-Cowan model in the stable focus regime. Analytical results provide a good approximation for response curves observed in patients with consistent significance. Additionally, we fitted the full non-linear Wilson-Cowan model to these patients, and we show that the model can fit in each case to the dynamics of patient tremor as well as the phase response curve, and the best fits are found to be stable foci for each patients (tied best fit in one instance). The model provides satisfactory prediction of how patient tremor will react to phase-locked stimulation by predicting patient amplitude response curves although they were not explicitly fitted. This can be partially explained by the relationship between the response curves in the model being compatible with what is found in the data. We also note that the non-linear Wilson-Cowan model is able to describe response to stimulation more precisely than the linearisation.
The fruit fly’s natural visual environment is often characterized by light intensities ranging across several orders of magnitude and by rapidly varying contrast across space and time. Fruit fly photoreceptors robustly transduce and, in conjunction with amacrine cells, process visual scenes and provide the resulting signal to downstream targets. Here, we model the first step of visual processing in the photoreceptor-amacrine cell layer. We propose a novel divisive normalization processor (DNP) for modeling the computation taking place in the photoreceptor-amacrine cell layer. The DNP explicitly models the photoreceptor feedforward and temporal feedback processing paths and the spatio-temporal feedback path of the amacrine cells. We then formally characterize the contrast gain control of the DNP and provide sparse identification algorithms that can efficiently identify each the feedforward and feedback DNP components. The algorithms presented here are the first demonstration of tractable and robust identification of the components of a divisive normalization processor. The sparse identification algorithms can be readily employed in experimental settings, and their effectiveness is demonstrated with several examples.
Morris-Lecar model is arguably the simplest dynamical model that retains both the slow-fast geometry of excitable phase portraits and the physiological interpretation of a conductance-based model. We augment this model with one slow inward current to capture the additional property of bistability between a resting state and a spiking limit cycle for a range of input current. The resulting dynamical system is a core structure for many dynamical phenomena such as slow spiking and bursting. We show how the proposed model combines physiological interpretation and mathematical tractability and we discuss the benefits of the proposed approach with respect to alternative models in the literature.
This is the first half of a two-part paper dealing with the geometry of color perception. Here we analyze in detail the seminal 1974 work by H.L. Resnikoff, who showed that there are only two possible geometric structures and Riemannian metrics on the perceived color space $\mathcal{P} $ compatible with the set of Schrödinger’s axioms completed with the hypothesis of homogeneity. We recast Resnikoff’s model into a more modern colorimetric setting, provide a much simpler proof of the main result of the original paper, and motivate the need of psychophysical experiments to confute or confirm the linearity of background transformations, which act transitively on $\mathcal{P} $. Finally, we show that the Riemannian metrics singled out by Resnikoff through an axiom on invariance under background transformations are not compatible with the crispening effect, thus motivating the need of further research about perceptual color metrics.
White matter pathways form a complex network of myelinated axons that regulate signal transmission in the nervous system and play a key role in behaviour and cognition. Recent evidence reveals that white matter networks are adaptive and that myelin remodels itself in an activity-dependent way, during both developmental stages and later on through behaviour and learning. As a result, axonal conduction delays continuously adjust in order to regulate the timing of neural signals propagating between different brain areas. This delay plasticity mechanism has yet to be integrated in computational neural models, where conduction delays are oftentimes constant or simply ignored. As a first approach to adaptive white matter remodeling, we modified the canonical Kuramoto model by enabling all connections with adaptive, phase-dependent delays. We analyzed the equilibria and stability of this system, and applied our results to two-oscillator and large-dimensional networks. Our joint mathematical and numerical analysis demonstrates that plastic delays act as a stabilizing mechanism promoting the network’s ability to maintain synchronous activity. Our work also shows that global synchronization is more resilient to perturbations and injury towards network architecture. Our results provide key insights about the analysis and potential significance of activity-dependent myelination in large-scale brain synchrony.
Neural populations with strong excitatory recurrent connections can support bistable states in their mean firing rates. Multiple fixed points in a network of such bistable units can be used to model memory retrieval and pattern separation. The stability of fixed points may change on a slower timescale than that of the dynamics due to short-term synaptic depression, leading to transitions between quasi-stable point attractor states in a sequence that depends on the history of stimuli. To better understand these behaviors, we study a minimal model, which characterizes multiple fixed points and transitions between them in response to stimuli with diverse time- and amplitude-dependencies. The interplay between the fast dynamics of firing rate and synaptic responses and the slower timescale of synaptic depression makes the neural activity sensitive to the amplitude and duration of square-pulse stimuli in a nontrivial, history-dependent manner. Weak cross-couplings further deform the basins of attraction for different fixed points into intricate shapes. We find that while short-term synaptic depression can reduce the total number of stable fixed points in a network, it tends to strongly increase the number of fixed points visited upon repetitions of fixed stimuli. Our analysis provides a natural explanation for the system's rich responses to stimuli of different durations and amplitudes while demonstrating the encoding capability of bistable neural populations for dynamical features of incoming stimuli.
