Adaptation of behavior requires the brain to change goals in a changing environment. Synaptic learning has demonstrated its effectiveness in changing the probability of selecting actions based on their outcome. In the extreme case, it is vital not to repeat an action to a given goal that led to harmful punishment. The present model proposes a simple neural mechanism of gain modulation that makes possible immediate changes in the probability of selecting a goal after punishment of variable intensity. Results show how gain modulation determine the type of elementary navigation process within the state space of a network of neuronal populations of excitatory neurons regulated by inhibition. Immediately after punishment, the system can avoid the punished populations by going back or by jumping to unpunished populations. This does not require particular credit assignment at the `choice' population but only gain modulation of neurons active at the time of punishment. Gain modulation does not require statistical relearning that may lead to further errors, but can encode memories of past experiences without modification of synaptic efficacies. Therefore, gain modulation can complements synaptic plasticity.
An important function of the brain is to predict which stimulus is likely to occur based on the perceived cues. The present research studied the branching behavior of a computational network model of populations of excitatory and inhibitory neurons, both analytically and through simulations. Results show how synaptic efficacy, retroactive inhibition and short-term synaptic depression determine the dynamics of selection between different branches predicting sequences of stimuli of different probabilities. Further results show that changes in the probability of the different predictions depend on variations of neuronal gain. Such variations allow the network to optimize the probability of its predictions to changing probabilities of the sequences without changing synaptic efficacy.
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.
In this article we present a biologically inspired model of activation of memory items in a sequence. Our model produces two types of sequences, corresponding to two different types of cerebral functions: activation of regular or irregular sequences. The switch between the two types of activation occurs through the modulation of biological parameters, without altering the connectivity matrix. Some of the parameters included in our model are neuronal gain, strength of inhibition, synaptic depression and noise. We investigate how these parameters enable the existence of sequences and influence the type of sequences observed. In particular we show that synaptic depression and noise drive the transitions from one memory item to the next and neuronal gain controls the switching between regular and irregular (random) activation.
Prediction is the ability of the brain to quickly activate a target concept in response to a related stimulus (prime). Experiments point to the existence of an overlap between the populations of the neurons coding for different stimuli, and other experiments show that primetarget relations arise in the process of long term memory formation. The classical modelling paradigm is that long term memories correspond to stable steady states of a Hopfield network with Hebbian connectivity. Experiments show that short term synaptic depression plays an important role in the processing of memories. This leads naturally to a computational model of priming, called latching dynamics; a stable state (prime) can become unstable and the system may converge to another transiently stable steady state (target). Hopfield network models of latching dynamics have been studied by means of numerical simulation, however the conditions for the existence of this dynamics have not been elucidated. In this work we use a combination of analytic and numerical approaches to confirm that latching dynamics can exist in the context of a symmetric Hebbian learning rule, however lacks robustness and imposes a number of biologically unrealistic restrictions on the model. In particular our work shows that the symmetry of the Hebbian rule is not an obstruction to the existence of latching dynamics, however fine tuning of the parameters of the model is needed.
We study pseudo-simple heteroclinic cycles for a Γ-equivariant system in R4 with finite Γ⊂O(4), and their nearby dynamics. In particular, in a first step towards a full classification – analogous to that which exists already for the class of simple cycles – we identify all finite subgroups of O(4) admitting pseudo-simple cycles. To this end we introduce a constructive method to build equivariant dynamical systems possessing a robust heteroclinic cycle. Extending a previous study we also investigate the existence of periodic orbits close to a pseudo-simple cycle, which depends on the symmetry groups of equilibria in the cycle. Moreover, we identify subgroups Γ⊂O(4), Γ⊄SO(4), admitting fragmentarily asymptotically stable pseudo-simple heteroclinic cycles. (It has been previously shown that for Γ⊂SO(4) pseudo-simple cycles generically are completely unstable.) Finally, we study a generalized heteroclinic cycle, which involves a pseudo-simple cycle as a subset.
Robust heteroclinic cycles in equivariant dynamical systems in \({\mathbb R}^4\) have been a subject of intense scientific investigation because, unlike heteroclinic cycles in \({\mathbb R}^3\), they can have an intricate geometric structure and complex asymptotic stability properties that are not yet completely understood. In a recent work, we have compiled an exhaustive list of finite subgroups of O(4) admitting the so-called simple heteroclinic cycles, and have identified a new class which we have called pseudo-simple heteroclinic cycles. By contrast with simple heteroclinic cycles, a pseudo-simple one has at least one equilibrium with an unstable manifold which has dimension 2 due to a symmetry. Here, we analyze the dynamics of nearby trajectories and asymptotic stability of pseudo-simple heteroclinic cycles in \({\mathbb R}^4\).
Prediction is the ability of the brain to quickly activate a target concept in response to a related stimulus (prime). Experiments point to the existence of an overlap between the populations of the neurons coding for different stimuli, and other experiments show that prime-target relations arise in the process of long term memory formation. The classical modelling paradigm is that long term memories correspond to stable steady states of a Hopfield network with Hebbian connectivity. Experiments show that short term synaptic depression plays an important role in the processing of memories. This leads naturally to a computational model of priming, called latching dynamics; a stable state (prime) can become unstable and the system may converge to another transiently stable steady state (target). Hopfield network models of latching dynamics have been studied by means of numerical simulation, however the conditions for the existence of this dynamics have not been elucidated. In this work we use a combination of analytic and numerical approaches to confirm that latching dynamics can exist in the context of a symmetric Hebbian learning rule, however lacks robustness and imposes a number of biologically unrealistic restrictions on the model. In particular our work shows that the symmetry of the Hebbian rule is not an obstruction to the existence of latching dynamics, however fine tuning of the parameters of the model is needed.
We review and extend the previous work where a model was introduced for Hopfield-type neural networks, which allows for the existence of heteroclinic dynamics between steady patterns. This dynamics is a mathematical model of periodic or aperiodic switching between stored information items in the brain, in particular, in the context of sequential memory or cognitive tasks as observed in experiments. The basic question addressed in this work is whether, given a sequence of steady patterns, it is possible by applying classical learning rules to build a matrix of connections between neurons in the network, such that a heteroclinic dynamics links these patterns. It has been shown previously that the answer is positive in the case where the sequence is a so-called simple consecutive cycle. We show that on the contrary the answer is negative for a non-simple cycle: heteroclinic dynamics does still exist; however, it cannot follow the sequence of patterns from which the connectivity matrix was derived.
This paper challenges and extends earlier seminal work. We consider the problem of describing mathematically the spontaneous activity of V1 by combining several important experimental observations including (1) the organization of the visual cortex into a spatially periodic network of hypercolumns structured around pinwheels, (2) the difference between short-range and long-range intracortical connections, the first ones being rather isotropic and producing naturally doubly periodic patterns by Turing mechanisms, the second one being patchy, and (3) the fact that the Turing patterns spontaneously produced by the short-range connections and the network of pinwheels have similar periods. By analyzing the PO maps, we are able to classify all possible singular points (the pinwheels) as having symmetries described by a small subset of the wallpaper groups. We then propose a description of the spontaneous activity of V1 using a classical voltage-based neural field model that features isotropic short-range connectivities modulated by non-isotropic long-range connectivities. A key observation is that, with only short-range connections and because the problem has full translational invariance in this case, a spontaneous doubly periodic pattern generates a 2-torus in a suitable functional space which persists as a flow-invariant manifold under small perturbations, for example when turning on the long-range connections. Through a complete analysis of the symmetries of the resulting neural field equation and motivated by a numerical investigation of the bifurcations of their solutions, we conclude that the branches of solutions which are stable over an extended range of parameters are those that correspond to patterns with an hexagonal (or nearly hexagonal) symmetry. The question of which patterns persist when turning on the long-range connections is answered by (1) analyzing the remaining symmetries on the perturbed torus and (2) combining this information with the Poincaré–Hopf theorem. We have developed a numerical implementation of the theory that has allowed us to produce the predicted patterns of activities, the planforms. In particular we generalize the contoured and non-contoured planforms predicted by previous authors.
It is widely believed that information is stored in the brain by means of the varying strength of synaptic connections between neurons. Stored patterns can be replayed upon the arrival of an appropriate stimulus. Hence, it is interesting to understand how an information pattern can be represented by the dynamics of the system. In this work, we consider a class of network neuron models, known as Hopfield networks, with a learning rule which consists of transforming an information string to a coupling pattern. Within this class of models, we study dynamic patterns, known as robust heteroclinic cycles, and establish a tight connection between their existence and the structure of the coupling.
In generic dynamical systems heteroclinic cycles are invariant sets of codimension at least one, but they can be structurally stable in systems which are equivariant under the action of a symmetry group, due to the existence of flow-invariant subspaces. For dynamical systems in R^n the minimal dimension for which such robust heteroclinic cycles can exist is n=3. In this case the list of admissible symmetry groups is short and well-known. The situation is different and more interesting when n=4. In this paper we list all finite groups Gamma such that an open set of smooth Gamma-equivariant dynamical systems in R^4 possess a very simple heteroclinic cycle (a structurally stable heteroclinic cycle satisfying certain additional constraints). This work extends the results which were obtained by Sottocornola in the case when all equilibria in the heteroclinic cycle belong to the same Gamma-orbit (in this case one speaks of homoclinic cycles).
The onset of convection for spherically invariant Rayleigh-B\'enard fluid flow is driven by marginal modes associated with spherical harmonics of a certain degree $\ell$, which depends upon the aspect ratio of the spherical shell. At certain critical values of the aspect ratio, marginal modes of degrees $\ell$ and $\ell+1$ coexist. Initially motivated by an experiment of electrophoretic convection between two concentric spheres carried in the International Space Station (GeoFlow project), we analyze the occurrence of intermittent dynamics near bifurcation in the case when marginal modes with $\ell=3, 4$ interact. The situation is by far more complex than in the well studied $\ell=1, 2$ mode interaction, however we show that heteroclinic cycles connecting equilibria with octahedral as well as axial symmetry can exist near bifurcation under certain conditions. Numerical simulations and continuation (using the software AUTO) on the center manifold help understanding these scenarios and show that the dynamics in these cases exhibits intermittent behaviour, even though the heteroclinic cycles may not be asymptotically stable in the usual sense.
In this paper we present an overview of pattern formation analysis for an analogue of the Swift-Hohenberg equation posed on the real hyperbolic space of dimension two, which we identify with the Poincaré disc \(\mathbb{D }\). Different types of patterns are considered: spatially periodic stationary solutions, radial solutions and traveling waves, however there are significant differences in the results with the Euclidean case. We apply equivariant bifurcation theory to the study of spatially periodic solutions on a given lattice of \(\mathbb{D }\) also called H-planforms in reference with the “planforms” introduced for pattern formation in Euclidean space. We consider in details the case of the regular octagonal lattice and give a complete descriptions of all H-planforms bifurcating in this case. For radial solutions (in geodesic polar coordinates), we present a result of existence for stationary localized radial solutions, which we have adapted from techniques on the Euclidean plane. Finally, we show that unlike the Euclidean case, the Swift-Hohenberg equation in the hyperbolic plane undergoes a Hopf bifurcation to traveling waves which are invariant along horocycles of \(\mathbb{D }\) and periodic in the “transverse” direction. We highlight our theoretical results with a selection of numerical simulations.
The existence of spatially localized solutions in neural networks is an important topic in neuroscience as these solutions are considered to characterize working (short-term) memory. We work with an unbounded neural network represented by the neural field equation with smooth firing rate function and a wizard hat spatial connectivity. Noting that stationary solutions of our neural field equation are equivalent to homoclinic orbits in a related fourth order ordinary differential equation, we apply normal form theory for a reversible Hopf bifurcation to prove the existence of localized solutions; further, we present results concerning their stability. Numerical continuation is used to compute branches of localized solution that exhibit snaking-type behaviour. We describe in terms of three parameters the exact regions for which localized solutions persist.
The primary visual cortex (V1) can be partitioned into fundamental domains or hypercolumns consisting of one set of orientation columns arranged around a singularity or ``pinwheel'' in the orientation preference map. A recent study on the specific problem of visual textures perception suggested that textures may be represented at the population level in the cortex as a second-order tensor, the structure tensor, within a hypercolumn. In this paper, we present a mathematical analysis of such interacting hypercolumns that takes into account the functional geometry of local and lateral connections. The geometry of the hypercolumn is identified with that of the Poincaré disk $\mathbb{D}$. Using the symmetry properties of the connections, we investigate the spontaneous formation of cortical activity patterns. These states are characterized by tuned responses in the feature space, which are doubly-periodically distributed across the cortex.
This paper completes the classification of bifurcation diagrams for H-planforms in the Poincaré disc \(\mathcal {D}\) whose fundamental domain is a regular octagon. An H-planform is a steady solution of a PDE or integro-differential equation in \(\mathcal {D}\), which is invariant under the action of a lattice subgroup Γ of U(1,1), the group of isometries of \({\mathcal{D}}\). In our case Γ generates a tiling of \(\mathcal {D}\) with regular octagons. This problem was introduced as an example of spontaneous pattern formation in a model of image feature detection by the visual cortex where the features are assumed to be represented in the space of structure tensors. Under ‘generic’ assumptions the bifurcation problem reduces to an ODE which is invariant by an irreducible representation of the group of automorphisms \(\mathcal {G}\) of the compact Riemann surface \(\mathcal {D}/\varGamma \). The irreducible representations of \(\mathcal {G}\) have dimensions one, two, three and four. The bifurcation diagrams for the representations of dimensions less than four have already been described and correspond to well-known group actions. In the present work we compute the bifurcation diagrams for the remaining three irreducible representations of dimension four, thus completing the classification. In one of these cases, there is generic bifurcation of a heteroclinic network connecting equilibria with two different orbit types.
Hopfield neural networks are simple models of information storage and retrieval, used both in the context of artificial intelligence and the brain [1]. Cycling orbits, that is periodic or recurrent solutions that visit multiple nodes of the network, are likely candidates for fragments of encoded information (working memory, cognitive tasks) [2, 3]. We conjecture that such cycles can be found near so-called robust heteroclinic cycles that exist for idealizations of a transformed version of the Hopfield network with voltage variables replaced by firing rate variables [4], [5]. We propose to conduct a numerical and theoretical study of robust heteroclinic cycles in the firing rate Hopfield networks and investigate their relation to the cycling orbits. A good starting point can be some work by Chuan Zhang available in the form of a preprint [6], [7]. The study requires basic knowledge of dynamical systems and their applications in computational biology. Some experience with MATLAB or a similar program is also an asset.
We study the neural field equations introduced by Chossat and Faugeras to model the representation and the processing of image edges and textures in the hypercolumns of the cortical area V1. The key entity, the structure tensor, intrinsically lives in a non-Euclidean, in effect hyperbolic, space. Its spatio-temporal behaviour is governed by nonlinear integro-differential equations defined on the Poincaré disc model of the two-dimensional hyperbolic space. Using methods from the theory of functional analysis we show the existence and uniqueness of a solution of these equations. In the case of stationary, that is, time independent, solutions we perform a stability analysis which yields important results on their behavior. We also present an original study, based on non-Euclidean, hyperbolic, analysis, of a spatially localised bump solution in a limiting case. We illustrate our theoretical results with numerical simulations. Mathematics Subject Classification:30F45, 33C05, 34A12, 34D20, 34D23, 34G20, 37M05, 43A85, 44A35, 45G10, 51M10, 92B20, 92C20.