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Journal of Mathematical Neuroscience logoLink to Journal of Mathematical Neuroscience
. 2011 Jun 6;1:4. doi: 10.1186/2190-8567-1-4

Analysis of a hyperbolic geometric model for visual texture perception

Gregory Faye 1,, Pascal Chossat 1,2, Olivier Faugeras 1
PMCID: PMC3280890  PMID: 22656402

Abstract

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, i.e. 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.

AMS subject classifications: 30F45, 33C05, 34A12, 34D20, 34D23, 34G20, 37M05, 43A85, 44A35, 45G10, 51M10, 92B20, 92C20.

Keywords: Neural fields, nonlinear integro-differential equations, functional analysis, non-Euclidean analysis, stability analysis, hyperbolic geometry, hypergeometric functions, bumps

1 Introduction

The selectivity of the responses of individual neurons to external features is often the basis of neuronal representations of the external world. For example, neurons in the primary visual cortex (V1) respond preferentially to visual stimuli that have a specific orientation [1-3], spatial frequency [4], velocity and direction of motion [5], color [6]. A local network in the primary visual cortex, roughly 1 mm2 of cortical surface, is assumed to consist of subgroups of inhibitory and excitatory neurons each of which is tuned to a particular feature of an external stimulus. These subgroups are the so-called Hubel and Wiesel hypercolumns of V1. We have introduced in [7] a new approach to model the processing of image edges and textures in the hypercolumns of area V1 that is based on a nonlinear representation of the image first order derivatives called the structure tensor [8,9]. We suggested that this structure tensor was represented by neuronal populations in the hypercolumns of V1. We also suggested that the time evolution of this representation was governed by equations similar to those proposed by Wilson and Cowan [10]. The question of whether some populations of neurons in V1 can represent the structure tensor is discussed in [7] but cannot be answered in a definite manner. Nevertheless, we hope that the predictions of the theory we are developing will help deciding on this issue.

Our present investigations were motivated by the work of Bressloff, Cowan, Golubitsky, Thomas and Wiener [11,12] on the spontaneous occurence of hallucinatory patterns under the influence of psychotropic drugs, and its extension to the structure tensor model. A further motivation was the following studies of Bressloff and Cowan [13,14,4] where they study a spatial extension of the ring model of orientation of Ben-Yishai [1] and Hansel, Sompolinsky [2]. To achieve this goal, we first have to better understand the local model, that is the model of a "texture" hypercolumn isolated from its neighbours.

The aim of this paper is to present a rigorous mathematical framework for the modeling of the representation of the structure tensor by neuronal populations in V1. We would also like to point out that the mathematical analysis we are developing here, is general and could be applied to other integro-differential equations defined on the set of structure tensors, so that even if the structure tensor were found to be not represented in a hypercolumn of V1, our framework would still be relevant. We then concentrate on the occurence of localized states, also called bumps. This is in contrast to the work of [7] and [15] where "spatially" periodic solutions were considered. The structure of this paper is as follows. In section 2 we introduce the structure tensor model and the corresponding equations. We also link our model to the ring model of orientations. In section 3 we use classical tools of evolution equations in functional spaces to analyse the problem of the existence and uniqueness of the solutions of our equations. In section 4 we study stationary solutions which are very important for the dynamics of the equation by analysing a nonlinear convolution operator and making use of the Haar measure of our feature space. In section 5, we push further the study of stationary solutions in a special case and we present a technical analysis involving hypergeometric functions of what we call a hyperbolic radially symmetric stationary-pulse in the high gain limit. Finally, in section 6, we present some numerical simulations of the solutions to verify the findings of the theoretical results.

2 The model

By definition, the structure tensor is based on the spatial derivatives of an image in a small area that can be thought of as part of a receptive field. These spatial derivatives are then summed nonlinearly over the receptive field. Let I(x, y) denote the original image intensity function, where x and y are two spatial coordinates. Let Inline graphic denote the scale-space representation of I obtained by convolution with the Gaussian kernel Inline graphic:

graphic file with name 2190-8567-1-4-i3.gif

The gradient Inline graphic is a two-dimensional vector of coordinates Inline graphic, Inline graphic which emphasizes image edges. One then forms the 2 × 2 symmetric matrix of rank one Inline graphic, where T indicates the transpose of a vector. The set of 2 × 2 symmetric positive semidefinite matrices of rank one will be noted S+(1, 2) throughout the paper (see [16] for a complete study of the set S+(p, n) of n × n symmetric positive semidefinite matrices of fixed-rank p <n). By convolving Inline graphic componentwise with a Gaussian Inline graphic we finally form the tensor structure as the symmetric matrix:

graphic file with name 2190-8567-1-4-i10.gif

where we have set for example:

graphic file with name 2190-8567-1-4-i11.gif

Since the computation of derivatives usually involves a stage of scale-space smoothing, the definition of the structure tensor requires two scale parameters. The first one, defined by σ1, is a local scale for smoothing prior to the computation of image derivatives. The structure tensor is insensitive to noise and details at scales smaller than σ1. The second one, defined by σ2, is an integration scale for accumulating the nonlinear operations on the derivatives into an integrated image descriptor. It is related to the characteristic size of the texture to be represented, and to the size of the receptive fields of the neurons that may represent the structure tensor.

By construction, Inline graphic is symmetric and non negative as Inline graphic by the inequality of Cauchy-Schwarz, then it has two orthonormal eigenvectors e1, e2 and two non negative corresponding eigenvalues λ1 and λ2 which we can always assume to be such that λ1 ≥ λ2 ≥ 0. Furthermore the spatial averaging distributes the information of the image over a neighborhood, and therefore the two eigenvalues are always positive. Thus, the set of the structure tensors lives in the set of 2 × 2 symmetric positive definite matrices, noted SPD(2, ℝ) throughout the paper. The distribution of these eigenvalues in the (λ1, λ2) plane reflects the local organization of the image intensity variations. Indeed, each structure tensor can be written as the linear combination:

graphic file with name 2190-8567-1-4-i14.gif (1)

where I2 is the identity matrix and Inline graphic. Some easy interpretations can be made for simple examples: constant areas are characterized by λ1 = λ2 ≈ 0, straight edges are such that λ1 ≫ λ2 ≈ 0, their orientation being that of e2, corners yield λ1 ≥ λ2 ≫ 0. The coherency c of the local image is measured by the ratio Inline graphic, large coherency reveals anisotropy in the texture.

We assume that a hypercolumn of V1 can represent the structure tensor in the receptive field of its neurons as the average membrane potential values of some of its membrane populations. Let Inline graphic be a structure tensor. The time evolution of the average potential Inline graphic for a given column is governed by the following neural mass equation adapted from [7] where we allow the connectivity function W to depend upon the time variable t and we integrate over the set of 2 × 2 symmetric definite-positive matrices:

graphic file with name 2190-8567-1-4-i18.gif (2)

The nonlinearity S is a sigmoidal function which may be expressed as:

graphic file with name 2190-8567-1-4-i19.gif

where μ describes the stiffness of the sigmoid. Iext is an external input.

The set SPD(2) can be seen as a foliated manifold by way of the set of special symmetric positive definite matrices SSPD(2) = SPD(2)∩SL(2, ℝ). Indeed, we have: Inline graphic. Furthermore, Inline graphic where Inline graphic is the Poincaré Disk, see e.g. [7]. As a consequence we use the following foliation of Inline graphic, which allows us to write for all Inline graphic, Inline graphic with Inline graphic. Inline graphic, z and Δ are related by the relation Inline graphic and the fact that z is the representation in Inline graphic of Inline graphic with Inline graphic.

It is well-known [17] that Inline graphic (and hence SSPD(2)) is a two-dimensional Riemannian space of constant sectional curvature equal to -1 for the distance noted d2 defined by

graphic file with name 2190-8567-1-4-i30.gif

The isometries of Inline graphic, that are the transformations that preserve the distance d2 are the elements of unitary group U(1, 1). In appendix A we describe the basic structure of this group. It follows, e.g. [18,7], that SDP(2) is a three-dimensional Riemannian space of constant sectional curvature equal to -1 for the distance noted d0 defined by

graphic file with name 2190-8567-1-4-i31.gif

As shown in proposition B.0.1 of appendix B it is possible to express the volume element Inline graphic in (z1, z2, Δ) coordinates with z = z1 + iz2:

graphic file with name 2190-8567-1-4-i33.gif

We note Inline graphic and equation (2) can be written in (z, Δ) coordinates:

graphic file with name 2190-8567-1-4-i35.gif

We get rid of the constant Inline graphic by redefining W as Inline graphic.

graphic file with name 2190-8567-1-4-i38.gif (3)

In [7], we have assumed that the representation of the local image orientations and textures is richer than, and contains, the local image orientations model which is conceptually equivalent to the direction of the local image intensity gradient. The richness of the structure tensor model has been expounded in [7]. The embedding of the ring model of orientation in the structure tensor model can be explained by the intrinsic relation that exists between the two sets of matrices SPD(2, ℝ) and S+(1, 2). First of all, when σ2 goes to zero, that is when the characteristic size of the structure becomes very small, we have Inline graphic, which means that the tensor Inline graphic degenerates to a tensor Inline graphic, which can be interpreted as the loss of one dimension. We can write each Inline graphic as Inline graphic, where u = (cos θ, sin θ)T and (r, θ) is the polar representation of x. Since, x and -x correspond to the same Inline graphic, θ is equated to θ + , Inline graphic. Thus Inline graphic, where ℙ1 is the real projective space of dimension 1 (lines of ℝ2). Then the integration scale σ2, at which the averages of the estimates of the image derivatives are computed, is the link between the classical representation of the local image orientations by the gradient and the representation of the local image textures by the structure tensor. It is also possible to highlight this explanation by coming back to the interpretation of straight edges of the previous paragraph. When λ1 ≫ λ2 ≈ 0 then Inline graphic and the orientation is that of e2. We denote by ℙ the projection of a 2 × 2 symmetric definite positive matrix on the set S+(1, 2) defined by:

graphic file with name 2190-8567-1-4-i46.gif

where Inline graphic is as in equation (1). We can introduce a metric on the set S+(1, 2) which is derived from a well-chosen Riemannian quotient geometry (see [16]). The resulting Riemannian space has strong geometrical properties: it is geodesically complete and the metric is invariant with respect to all transformations that preserve angles (orthogonal transformations, scalings and pseudoinversions). Related to the decomposition Inline graphic, a metric on the space S+(1, 2) is given by:

graphic file with name 2190-8567-1-4-i47.gif

The space S+(1, 2) endowed with this metric is a Riemannian manifold (see [16]). Finally, the distance associated to this metric is given by:

graphic file with name 2190-8567-1-4-i48.gif

where Inline graphic and (ri, θi) denotes the polar coordinates of xi for i = 1, 2. The volume element in (r, θ) coordinates is:

graphic file with name 2190-8567-1-4-i50.gif

where we normalize to 1 the volume element for the θ coordinate.

Let now Inline graphic be a symmetric positive semidefinite matrix. The average potential V(τ, t) of the column has its time evolution that is governed by the following neural mass equation which is just a projection of equation (2) on the subspace S+(1, 2):

graphic file with name 2190-8567-1-4-i52.gif (4)

In (r, θ) coordinates, (4) is rewritten as:

graphic file with name 2190-8567-1-4-i53.gif

This equation is richer than the ring model of orientation as it contains an additional information on the contrast of the image in the orthogonal direction of the prefered orientation. If one wants to recover the ring model of orientation tuning in the visual cortex as it has been presented and studied by [1,2,19], it is sufficient i) to assume that the connectivity function is time-independent and has a convolutional form:

graphic file with name 2190-8567-1-4-i54.gif

and ii) to look at semi-homogeneous solutions of equation 4, i.e., solutions which do not depend upon the variable r. We finally obtain:

graphic file with name 2190-8567-1-4-i55.gif (5)

where:

graphic file with name 2190-8567-1-4-i56.gif

It follows from the above discussion that the structure tensor contains, at a given scale, more information than the local image intensity gradient at the same scale and that it is possible to recover the ring model of orientations from the structure tensor model.

The aim of the following sections is to establish that (3) is well-defined and to give necessary and sufficient conditions on the different parameters in order to prove some results on the existence and uniqueness of a solution of (3).

3 The existence and uniqueness of a solution

In this section we provide theoretical and general results of existence and uniqueness of a solution of (2). In the first subsection 3.1 we study the simpler case of the homogeneous solutions of (2), i.e. of the solutions that are independent of the tensor variable Inline graphic. This simplified model allows us to introduce some notations for the general case and to establish the useful lemma 3.1.1. We then prove in 3.2 the main result of this section, that is the existence and uniqueness of a solution of (2). Finally we develop the useful case of the semi-homogeneous solutions of (2), i.e. of solutions that depend on the tensor variable but only through its z coordinate in Inline graphic.

3.1 Homogeneous solutions

A homogeneous solution to (2) is a solution V that does not depend upon the tensor variable Inline graphic for a given homogenous input I(t) and a constant initial condition V0. In (z, Δ) coordinates, a homogeneous solution of (3) is defined by:

graphic file with name 2190-8567-1-4-i57.gif

where:

graphic file with name 2190-8567-1-4-i58.gif (6)

Hence necessary conditions for the existence of a homogeneous solution are that:

• the double integral (6) is convergent,

Inline graphic does not depend upon the variable (z, Δ). In that case, we write Inline graphic instead of Inline graphic.

In the special case where W(z, Δ, z', Δ', t) is a function of only the distance d0 between (z, Δ) and (z', Δ'):

graphic file with name 2190-8567-1-4-i61.gif

the second condition is automatically satisfied. The proof of this fact is given in lemma D.0.2 of appendix D. To summarize, the homogeneous solutions satisfy the differential equation:

graphic file with name 2190-8567-1-4-i62.gif (7)

3.1.1 A first existence and uniqueness result

Equation (3) defines a Cauchy's problem and we have the following theorem.

Theorem 3.1.1. If the external input Iext(t) and the connectivity function Inline graphicare continuous on some closed interval J containing 0, then for all V0 in ℝ, there exists a unique solution of (7) defined on a subinterval J0 of J containing 0 such that V (0) = V0.

Proof. It is a direct application of Cauchy's theorem on differential equations. We consider the mapping f : J × ℝ → ℝ defined by:

graphic file with name 2190-8567-1-4-i63.gif

It is clear that f is continuous from J × ℝ to ℝ. We have for all x, y ∈ ℝ and t J:

graphic file with name 2190-8567-1-4-i64.gif

where Inline graphic.

Since, Inline graphic is continuous on the compact interval J, it is bounded there by C > 0 and:

graphic file with name 2190-8567-1-4-i67.gif

We can extend this result to the whole time real line if I and Inline graphic are continuous on ℝ.

Proposition 3.1.1. If Iext and Inline graphicare continuous on +, then for all V0 in , there exists a unique solution of (7) defined on + such that V (0) = V0.

Proof. We have already shown the following inequality:

graphic file with name 2190-8567-1-4-i68.gif

Then f is locally Lipschitz with respect to its second argument. Let V be a maximal solution on J0 and we denote by β the upper bound of J0. We suppose that β < + ∞. Then we have for all t ≥ 0:

graphic file with name 2190-8567-1-4-i69.gif

where Sm = supx∈ℝ |S(x)|.

This implies that the maximal solution V is bounded for all t ∈ [0, β], but theorem C.0.2 of appendix C ensures that it is impossible. Then, it follows that necessarily β = + ∞.   □

3.1.2 Simplification of (6) in a special case

Invariance

In the previous section, we have stated that in the special case where W was a function of the distance between two points in Inline graphic, then Inline graphic did not depend upon the variables (z, Δ). As already said in the previous section, the following result holds (see proof of lemma D.0.2 of appendix D).

Lemma 3.1.1. Suppose that W is a function of d0 Inline graphic only. Then Inline graphicdoes not depend upon the variable Inline graphic.

Mexican hat connectivity

In this paragraph, we push further the computation of Inline graphic in the special case where W does not depend upon the time variable t and takes the special form suggested by Amari in [20], commonly referred to as the "Mexican hat" connectivity. It features center excitation and surround inhibition which is an effective model for a mixed population of interacting inhibitory and excitatory neurons with typical cortical connections. It is also only a function of d0 Inline graphic.

In detail, we have:

graphic file with name 2190-8567-1-4-i72.gif

where:

graphic file with name 2190-8567-1-4-i73.gif

with 0 ≤ σ1 σ2 and 0 ≤ A ≤ 1.

In this case we can obtain a very simple closed-form formula for Inline graphic as shown in the following lemma.

Lemma 3.1.2. When W is the specific Mexican hat function just defined then:

graphic file with name 2190-8567-1-4-i74.gif (8)

where erf is the error function defined as:

graphic file with name 2190-8567-1-4-i75.gif

Proof. The proof is given in lemma E.0.3 of appendix E.   □

3.2 General solution

We now present the main result of this section about the existence and uniqueness of solutions of equation (2). We first introduce some hypotheses on the connectivity function W. We present them in two ways: first on the set of structure tensors considered as the set SPD(2), then on the set of tensors seen as Inline graphic. Let J be a subinterval of ℝ. We assume that:

Inline graphic,

Inline graphic where W is defined as Inline graphic for all Inline graphic where Id2 is the identity matrix of ℳ2(ℝ),

Inline graphic where Inline graphic.

Equivalently, we can express these hypotheses in (z, Δ) coordinates:

Inline graphic,

Inline graphic where W is defined as W(z, Δ, t) = W (d2(z, 0), | log(Δ)|, t) for all Inline graphic,

Inline graphic where

graphic file with name 2190-8567-1-4-i87.gif

3.2.1 Functional space setting

We introduce the following mapping fg : (t, ϕ) → fg(t, ϕ) such that:

graphic file with name 2190-8567-1-4-i88.gif (9)

Our aim is to find a functional space Inline graphic where (3) is well-defined and the function fg maps Inline graphic to Inline graphic for all ts. A natural choice would be to choose ϕ as a Inline graphic-integrable function of the space variable with 1 ≤ p < +∞. Unfortunately, the homogeneous solutions (constant with respect to (z, Δ)) do not belong to that space. Moreover, a valid model of neural networks should only produce bounded membrane potentials. That is why we focus our choice on the functional space Inline graphic. As Inline graphic is an open set of ℝ3, Inline graphic is a Banach space for the norm:Inline graphic.

Proposition 3.2.1. If Inline graphic with Inline graphic and W satisfies hypotheses (H1bis)-(H3bis) then f g is well-defined and is from J × Inline graphic to Inline graphic.

Proof. Inline graphic, we have:

graphic file with name 2190-8567-1-4-i96.gif

   □

3.2.2 The existence and uniqueness of a solution of (3)

We rewrite (3) as a Cauchy problem:

graphic file with name 2190-8567-1-4-i97.gif (10)

Theorem 3.2.1. If the external current Iext belongs to Inline graphic with J an open interval containing 0 and W satisfies hypotheses (H1bis)-(H3bis), then for all V0 Inline graphic, there exists a unique solution of (10) defined on a subinterval J0 of J containing 0 such that V (z, Δ, 0) = V0(z, Δ) for all Inline graphic.

Proof. We prove that fg is continuous on J × Inline graphic. We have

graphic file with name 2190-8567-1-4-i100.gif

and therefore

graphic file with name 2190-8567-1-4-i101.gif

Because of condition (H2) we can choose |t -s| small enough so that Inline graphic is arbitrarily small. This proves the continuity of fg. Moreover it follows from the previous inequality that:

graphic file with name 2190-8567-1-4-i103.gif

with Inline graphic. This ensures the Lipschitz continuity of fg with respect to its second argument, uniformly with respect to the first. The Cauchy-Lipschitz theorem on a Banach space yields the conclusion.   □

Remark 3.2.1. Our result is quite similar to those obtained by Potthast and Graben in [21]. The main differences are that, first, we allow the connectivity function to depend upon the time variable t and, second, that our space features is no longer a n but a Riemanian manifold. In their article, Potthast and Graben also work with a different functional space by assuming more regularity for the connectivity function W and then obtain more regularity for their solutions.

Proposition 3.2.2. If the external current Iext belongs to Inline graphic and W satisfies hypotheses (H1bis)-(H3bis) with J = +, then for all V0 Inline graphic, there exists a unique solution of (10) defined on + such that V (z, Δ, 0) = V0(z, Δ) for all Inline graphic.

Proof. We have just seen in the previous proof that fg is globally Lipschitz with respect to its second argument:

graphic file with name 2190-8567-1-4-i107.gif

then theorem C.0.3 of the appendix C gives the conclusion.   □

3.2.3 The intrinsic boundedness of a solution of (3)

In the same way as in the homogeneous case, we show a result on the boundedness of a solution of (3).

Proposition 3.2.3. If the external current Iext belongs to Inline graphicand is bounded in time Inline graphic and W satisfies hypotheses (H1bis)-(H3bis) with J = +, then the solution of (10) is bounded for each initial condition V0 Inline graphic.

Let us set:

graphic file with name 2190-8567-1-4-i109.gif

where Inline graphic.

Proof. Let V be a solution defined on ℝ+. Then we have for all t ∈ ℝ+*:

graphic file with name 2190-8567-1-4-i111.gif

The following upperbound holds

graphic file with name 2190-8567-1-4-i112.gif (11)

We can rewrite (11) as:

graphic file with name 2190-8567-1-4-i113.gif (12)

If Inline graphic this implies Inline graphic for all t > 0 and hence ||V(t)||<ρg for all t > 0, proving that Bp is stable. Now assume that ||V(t)||<ρg for all t ≥ 0. The inequality (12) shows that for t large enough this yields a contradiction. Therefore there exists t0 > 0 such that ||V(t0)||<ρg. At this time instant we have

graphic file with name 2190-8567-1-4-i116.gif

and hence

graphic file with name 2190-8567-1-4-i117.gif

   □

The following corollary is a consequence of the previous proposition.

Corollary 3.2.1. If Inline graphic and Inline graphic then:

graphic file with name 2190-8567-1-4-i120.gif

3.3 Semi-homogeneous solutions

A semi-homogeneous solution of (3) is defined as a solution which does not depend upon the variable Δ. In other words, the populations of neurons is not sensitive to the determinant of the structure tensor, that is to the contrast of the image intensity. The neural mass equation is then equivalent to the neural mass equation for tensors of unit determinant. We point out that semi-homogeneous solutions were previously introduced in [7] where a bifurcation analysis of what they called H-planforms was performed. In this section, we define the framework in which their equations make sense without giving any proofs of our results as it is a direct consequence of those proven in the general case. We rewrite equation (3) in the case of semi-homogeneous solutions:

graphic file with name 2190-8567-1-4-i121.gif (13)

where

graphic file with name 2190-8567-1-4-i122.gif

We have implicitly made the assumption, that Wsh does not depend on the coordinate Δ. Some conditions under which this assumption is satisfied are described below and are the direct transductions of those of the general case in the context of semi-homogeneous solutions.

Let J be an open interval of ℝ. We assume that:

Inline graphic,

Inline graphic where Wsh is defined as Wsh (z, t) = wsh(d2(z, 0), t) for all Inline graphic,

Inline graphic where Inline graphic.

Note that conditions (C1)-(C2) and lemma 3.1.1 imply that for all Inline graphic, Inline graphic. And then, for all Inline graphic, the mapping z' → Wsh(z, z', t) is integrable on Inline graphic.

From now on, Inline graphic and the Fischer-Riesz's theorem ensures that Inline graphic is a Banach space for the norm: Inline graphic.

Theorem 3.3.1. If the external current Iext belongs to Inline graphicwith J an open interval containing 0 and Wsh satisfies conditions (C1)-(C3), then for all V0 Inline graphic, there exists a unique solution of (13) defined on a subinterval J0 of J containing 0.

This solution, defined on the subinterval J of ℝ can in fact be extended to the whole real line, and we have the following proposition.

Proposition 3.3.1. If the external current Iext belongs to Inline graphicand Wsh satisfies conditions (C1)-(C3) with J = +, then for all V0 Inline graphic, there exists a unique solution of (13) defined on +.

We can also state a result on the boundedness of a solution of (13):

Proposition 3.3.2. Let Inline graphic, with Inline graphic. The open ball Bp of Inline graphic of center 0 and radius p is stable under the dynamics of equation (13). Moreover it is an attracting set for this dynamics and if V0 Bρ and T = inf{t > 0 such that V(t) ∈ BP} then:

graphic file with name 2190-8567-1-4-i135.gif

4 Stationary solutions

We look at the equilibrium states, noted Inline graphic of (3), when the external input I and the connectivity W do not depend upon the time. We assume that W satisfies hypotheses (H1bis)-(H2bis). We redefine for convenience the sigmoidal function to be:

graphic file with name 2190-8567-1-4-i137.gif

so that a stationary solution (independent of time) satisfies:

graphic file with name 2190-8567-1-4-i138.gif (14)

We define the nonlinear operator from Inline graphic to Inline graphic, noted Inline graphic, by:

graphic file with name 2190-8567-1-4-i140.gif (15)

Finally, (14) is equivalent to:

graphic file with name 2190-8567-1-4-i141.gif

4.1 Study of the nonlinear operator Inline graphic

We recall that we have set for the Banach space Inline graphic and proposition 3.2.1 shows that Inline graphic. We have the further properties:

Proposition 4.1.1. Inline graphicsatisfies the following properties:

Inline graphic,

Inline graphicis continuous on +,

Proof. The first property was shown to be true in the proof of theorem 3.3.1. The second property follows from the following inequality:

graphic file with name 2190-8567-1-4-i146.gif

We denote by Inline graphic and Inline graphic the two operators from Inline graphic to Inline graphic defined as follows for all V Inline graphic and all Inline graphic:

graphic file with name 2190-8567-1-4-i150.gif (16)

and

graphic file with name 2190-8567-1-4-i151.gif

where H is the Heaviside function.

It is straightforward to show that both operators are well-defined on Inline graphic and map Inline graphic to Inline graphic. Moreover the following proposition holds.

Proposition 4.1.2. We have

graphic file with name 2190-8567-1-4-i152.gif

Proof. It is a direct application of the dominated convergence theorem using the fact that:

graphic file with name 2190-8567-1-4-i153.gif

   □

4.2 The convolution form of the operator Inline graphicin the semi-homogeneous case

It is convenient to consider the functional space Inline graphic to discuss semi-homogeneous solutions. A semi-homogeneous persistent state of (3) is deduced from (14) and satisfies:

graphic file with name 2190-8567-1-4-i155.gif (17)

where the nonlinear operator Inline graphic from Inline graphicsh to Inline graphicsh is defined for all V ∈ Inline graphicshand Inline graphic; by:

graphic file with name 2190-8567-1-4-i157.gif

We define the associated operators, Inline graphic, Inline graphic:

graphic file with name 2190-8567-1-4-i160.gif
graphic file with name 2190-8567-1-4-i161.gif

We rewrite the operator Inline graphic in a convenient form by using the convolution in the hyperbolic disk. First, we define the convolution in a such space. Let O denote the center of the Poincaré disk that is the point represented by z = 0 and dg denote the Haar measure on the group G = SU(1, 1) (see [22] and appendix A), normalized by:

graphic file with name 2190-8567-1-4-i162.gif

for all functions of Inline graphic. Given two functions f1, f2 in Inline graphic we define the convolution * by:

graphic file with name 2190-8567-1-4-i164.gif

We recall the notation Inline graphic.

Proposition 4.2.1. For all μ ≥ 0 and V Inline graphicsh we have:

graphic file with name 2190-8567-1-4-i166.gif (18)

Proof. We only prove the result forInline graphic. Let Inline graphic, then:

graphic file with name 2190-8567-1-4-i167.gif

and for all g SU(1, 1), d2(z, z') = d2(g·z, g·z') so that:

graphic file with name 2190-8567-1-4-i168.gif

   □

Let b be a point on the circle Inline graphic. For Inline graphic, we define the "inner product" <z, b > to be the algebraic distance to the origin of the (unique) horocycle based at b through z (see [7]). Note that <z, b > does not depend on the position of z on the horocycle. The Fourier transform in Inline graphic is defined as (see [22]):

graphic file with name 2190-8567-1-4-i170.gif

for a function Inline graphic such that this integral is well-defined.

Lemma 4.2.1. The Fourier transform in Inline graphic, Inline graphic of Wsh does not depend upon the variable Inline graphic. Proof. For all λ ∈ ℝ and Inline graphicInline graphic,

graphic file with name 2190-8567-1-4-i176.gif

We recall that for all ϕ ∈ ℝ rϕ is the rotation of angle ϕ and we have Wsh(rϕ ·z) = Wsh(z), dm(z) = dm(rϕ ·z) and <z, b > = <rϕ ·z, rϕ ·b >, then:

graphic file with name 2190-8567-1-4-i177.gif

   □

We now introduce two functions that enjoy some nice properties with respect to the Hyperbolic Fourier transform and are eigenfunctions of the linear operator Inline graphic.

Proposition 4.2.2. Let eλ, b(z) = e(-+1)<z, b>and Inline graphic then:

Inline graphic

Inline graphic

Proof. We begin with Inline graphic and use the horocyclic coordinates. We use the same changes of variables as in lemma 3.1.1:

graphic file with name 2190-8567-1-4-i182.gif

By rotation, we obtain the property for all Inline graphic.

For the second property [[22], Lemma 4.7] shows that:

graphic file with name 2190-8567-1-4-i183.gif

A consequence of this proposition is the following lemma.   □

Lemma 4.2.2. The linear operator Inline graphicis not compact and for all μ ≥ 0, the nonlinear operator Inline graphicis not compact.

Proof. The previous proposition 4.2.2 shows that Inline graphic has a continuous spectrum which iimplies that is not a compact operator.

Let U be in Inline graphicsh, for all V Inline graphicsh we differentiate Inline graphic and compute its Frechet derivative:

graphic file with name 2190-8567-1-4-i184.gif

If we assume further that U does not depend upon the space variable z, U(z) = U0 we obtain:

graphic file with name 2190-8567-1-4-i185.gif

If Inline graphic was a compact operator then its Frechet derivative Inline graphic would also be a compact operator, but it is impossible. As a consequence, Inline graphic is not a compact operator.   □

4.3 The convolution form of the operator Inline graphicin the general case

We adapt the ideas presented in the previous section in order to deal with the general case. We recall that if H is the group of positive real numbers with multiplication as operation, then the Haar measure dh is given by Inline graphic. For two functions f1, f2 in Inline graphic we define the convolution ⋆ by:

graphic file with name 2190-8567-1-4-i189.gif

We recall that we have set by definition: W(z, Δ) = W(d2(z, 0), |log(Δ)|).

Proposition 4.3.1. For all μ ≥ 0 and V Inline graphicwe have:

graphic file with name 2190-8567-1-4-i190.gif (19)

Proof. Let (z, Δ) be in Inline graphic. We follow the same ideas as in proposition 4.2.1 and prove only the first result. We have

graphic file with name 2190-8567-1-4-i192.gif

   □

We next assume further that the function W is separable in z, Δ and more precisely that W(z, Δ) = W1(z)W2(log(Δ)) where W1(z) = W1(d2(z, 0)) and W2(log(Δ)) = W2(|log(Δ)|) for all Inline graphic. The following proposition is an echo to proposition 4.2.2.

Proposition 4.3.2. Let eλ,b(z) = e(-+1)<z, b>, Inline graphic and hξ(Δ) = elog(Δ) then:

Inline graphic

Inline graphic

Where Inline graphic is the usual Fourier transform of W2.

Proof. The proof of this proposition is exactly the same as for proposition 4.2.2. Indeed:

graphic file with name 2190-8567-1-4-i198.gif

   □

A straightforward consequence of this proposition is an extension of lemma 4.2.2 to the general case:

Lemma 4.3.1. The linear operator Inline graphicis not compact and for all μ ≥ 0, the nonlinear operator Inline graphicis not compact.

4.4 The set of the solutions of (14)

Let Inline graphic be the set of the solutions of (14) for a given slope parameter μ:

graphic file with name 2190-8567-1-4-i199.gif

We have the following proposition.

Proposition 4.4.1. If the input current Iext is equal to a constant Inline graphic, i.e. does not depend upon the variables (z, Δ) then for all Inline graphic, Inline graphic. In the general case Inline graphic, if the condition Inline graphicis satisfied, then Card Inline graphic.

Proof. Due to the properties of the sigmoid function, there always exists a constant solution in the case where Iext is constant. In the general case where Inline graphic, the statement is a direct application of the Banach fixed point theorem, as in [23].   □

Remark 4.4.1. If the external input does not depend upon the variables (z, Δ) and if the condition Inline graphicis satisfied, then there exists a unique stationary solution by application of proposition 4.4.1. Moreover, this stationary solution does not depend upon the variables (z, Δ) because there always exists one constant stationary solution when the external input does not depend upon the variables (z, Δ). Indeed equation (14) is then equivalent to:

graphic file with name 2190-8567-1-4-i206.gif

and β does not depend upon the variables (z, Δ) because of lemma 3.1.1. Because of the property of the sigmoid function S, the equation Inline graphic has always one solution.

If on the other hand the input current does depend upon these variables, is invariant under the action of a subgroup of U(1, 1), the group of the isometries of Inline graphic(see appendix A), and the condition Inline graphicis satisfied, then the unique stationary solution will also be invariant under the action of the same subgroup. We refer the interested reader to our work [15] on equivariant bifurcation of hyperbolic planforms on the subject.

When the condition Inline graphicis satisfied we call primary stationary solution the unique solution in Inline graphic.

4.5 Stability of the primary stationary solution

In this subsection we show that the condition Inline graphic guarantees the stability of the primary stationary solution to (3).

Theorem 4.5.1. We suppose that Inline graphic and that the condition Inline graphicis satisfied, then the associated primary stationary solution of (3) is asymtotically stable.

Proof. Let Inline graphic be the primary stationary solution of (3), as Inline graphic is satisfied. Let also Vμ be the unique solution of the same equation with some initial condition Inline graphic, see theorem 3.3.1. We introduce a new function Inline graphic which satisfies:

graphic file with name 2190-8567-1-4-i213.gif

where Inline graphic, Inline graphic and the vector Θ(X(z, Δ, t)) is given by Inline graphic with Inline graphic. We note that, because of the definition of Θ and the mean value theorem |Θ(X(z, Δ, t))| ≤ μ|X(z, Δ, t)|. This implies that |Θ(r)| ≤ |r| for all r ∈ ℝ.

graphic file with name 2190-8567-1-4-i218.gif

If we set: G(t) = eαt||X(t)||, then we have:

graphic file with name 2190-8567-1-4-i219.gif

and G is continuous for all t ≥ 0. The Gronwall inequality implies that:

graphic file with name 2190-8567-1-4-i220.gif

and the conclusion follows.   □

5 Spatially localised bumps in the high gain limit

In many models of working memory, transient stimuli are encoded by feature-selective persistent neural activity. Such stimuli are imagined to induce the formation of a spatially localised bump of persistent activity which coexists with a stable uniform state. As an example, Camperi and Wang [24] have proposed and studied a network model of visuo-spatial working memory in prefontal cortex adapted from the ring model of orientation of Ben-Yishai and colleagues [1]. Many studies have emerged in the past decades to analyse these localised bumps of activity [25-29], see the paper by Coombes for a review of the domain [30]. In [25,26,28], the authors have examined the existence and stability of bumps and multi-bumps solutions to an integro-differential equation describing neuronal activity along a single spatial domain. In [27,29] the study is focused on the two-dimensional model and a method is developed to approximate the integro-differential equation by a partial differential equation which makes possible the determination of the stability of circularly symmetric solutions. It is therefore natural to study the emergence of spatially localized bumps for the structure tensor model in a hypercolumn of V1. We only deal with the reduced case of equation (13) which means that the membrane activity does not depend upon the contrast of the image intensity, keeping the general case for future work.

In order to construct exact bump solutions and to compare our results to previous studies [25-29], we consider the high gain limit μ → ∞ of the sigmoid function. As above we denote by H the Heaviside function defined by H(x) = 1 for x ≥ 0 and H(x) = 0 otherwise. Equation (13) is rewritten as:

graphic file with name 2190-8567-1-4-i221.gif (20)

We have introduced a threshold κ to shift the zero of the Heaviside function. We make the assumption that the system is spatially homogeneous that is, the external input I does not depend upon the variables t and the connectivity function depends only on the hyperbolic distance between two points of Inline graphic. For illustrative purposes, we will use the exponential weight distribution as a specific example throughout this section:

graphic file with name 2190-8567-1-4-i223.gif (21)

The theoretical study of equation (20) has been done in [21] where the authors have imposed strong regularity assumptions on the kernel function W, such as Hölder continuity, and used compactness arguments and integral equation techniques to obtain a global existence result of solutions to (20). Our approach is very different, we follow that of [25,31,29] by proceeding in a constructive fashion. In a first part, we define what we call a hyperbolic radially symmetric bump and present some preliminary results for the linear stability analysis of the last part. The second part is devoted to the proof of a technical theorem 5.1.1 which is stated in the first part. The proof uses results on the Fourier transform introduced in section 4, hyperbolic geometry and hypergeometric functions. Our results will be illustrated in the following section 6.

5.1 Existence of hyperbolic radially symmetric bumps

From equation (20) a general stationary pulse satisfies the equation:

graphic file with name 2190-8567-1-4-i224.gif

For convenience, we note M(z, K) the integral K W(z, z')dm(z') with Inline graphic. The relation V (z) = κ holds for all z ∈ ∂K.

Definition 5.1.1. V is called a hyperbolic radially symmetric stationary-pulse solution of (20) if V depends only upon the variable r and is such that:

graphic file with name 2190-8567-1-4-i226.gif

and is a fixed point of equation (20):

graphic file with name 2190-8567-1-4-i227.gif (22)

where Inline graphic is a Gaussian input and Inline graphic is defined by the following equation:

graphic file with name 2190-8567-1-4-i230.gif

and Bh(0, ω) is a hyperbolic disk centered at the origin of hyperbolic radius ω.

From symmetry arguments there exists a hyperbolic radially symmetric stationary-pulse solution V(r) of (20), furthermore the threshold κ and width ω are related according to the self-consistency condition

graphic file with name 2190-8567-1-4-i231.gif (23)

where

graphic file with name 2190-8567-1-4-i232.gif

The existence of such a bump can then be established by finding solutions to (23) The function N(ω) is plotted in Figure 1 for a range of the input amplitude Inline graphic. The horizontal dashed lines indicate different values of ακ, the points of intersection determine the existence of stationary pulse solutions. Qualitatively, for sufficiently large input amplitude Inline graphic we have N'(0) < 0 and it is possible to find only one solution branch for large ακ. For small input amplitudes Inline graphic we have N'(0) > 0 and there always exists one solution branch for αβ <γc ≈ 0.06. For intermediate values of the input amplitude Inline graphic, as αβ varies, we have the possiblity of zero, one or two solutions. Anticipating the stability results of section 5.3, we obtain that when N'(ω) < 0 then the corresponding solution is stable.

Figure 1.

Figure 1

Plot of N(ω) defined in (23) as a function of the pulse width ω for several values of the input amplitude Inline graphic and for a fixed input width σ = 0.05. The horizontal dashed lines indicate different values of ακ. The connectivity function is given in equation (21) and the parameter b is set to b = 0.2.

We end this subsection with the usefull and technical following formula.

Theorem 5.1.1. For all Inline graphic:

graphic file with name 2190-8567-1-4-i235.gif (24)

Where Inline graphic is the Fourier Helgason transform of Inline graphic and

graphic file with name 2190-8567-1-4-i238.gif

with α + β + 1 = ρ and F is the hypergeometric function of first kind.

Remark 5.1.1. We recall that F admits the integral representation [32]:

graphic file with name 2190-8567-1-4-i239.gif

with ℜ(γ) > ℜ(β) > 0.

Remark 5.1.2. In section 4 we introduced the function Inline graphic. In [22], it is shown that:

graphic file with name 2190-8567-1-4-i241.gif

Remark 5.1.3. Let us point out that this result can be linked to the work of Folias and Bressloff in [31] and then used in [29]. They constructed a two-dimensional pulse for a general, radially symmetric synaptic weight function. They obtain a similar formal representation of the integral of the connectivity function w over the disk B(O, a) centered at the origin O and of radius a. Using their notations,

graphic file with name 2190-8567-1-4-i242.gif

where Jν(x) is the Bessel function of the first kind and Inline graphic is the real Fourier transform of w. In our case, instead of the Bessel function, we find Inline graphic which is linked to the hypergeometric function of the first kind.

We now show that for a general monotonically decreasing weight function W, the function Inline graphic is necessarily a monotonically decreasing function of r. This will ensure that the hyperbolic radially symmetric stationary-pulse solution (22) is also a monotonically decreasing function of r in the case of a Gaussian input. The demonstration of this result will directly use theorem 5.1.1.

Proposition 5.1.1. V is a monotonically decreasing function in r for any monotonically decreasing synaptic weight function W.

Proof. Differentiating ℳ with respect to r yields:

graphic file with name 2190-8567-1-4-i245.gif

We have to compute

graphic file with name 2190-8567-1-4-i246.gif

It is result of elementary hyperbolic trigonometry that

graphic file with name 2190-8567-1-4-i247.gif (25)

we let ρ = tanh(r), ρ' = tanh(r') and define

graphic file with name 2190-8567-1-4-i248.gif

It follows that

graphic file with name 2190-8567-1-4-i249.gif

and

graphic file with name 2190-8567-1-4-i250.gif

We conclude that if ρ > tanh(ω) then for all 0 ≤ ρ' ≤ tanh(ω) and 0 ≤ θ ≤ 2π

graphic file with name 2190-8567-1-4-i251.gif

which implies Inline graphic for r >ω, since W' < 0.

To see that it is also negative for r <ω, we differentiate equation (24) with respect to r:

graphic file with name 2190-8567-1-4-i253.gif

The following formula holds for the hypergeometric function (see Erdelyi in [32]):

graphic file with name 2190-8567-1-4-i254.gif

It implies

graphic file with name 2190-8567-1-4-i255.gif

Substituting in the previous equation giving Inline graphic we find:

graphic file with name 2190-8567-1-4-i257.gif

implying that:

graphic file with name 2190-8567-1-4-i258.gif

Consequently, Inline graphic for r <ω. Hence V is monotonically decreasing in r for any monotonically decreasing synaptic weight function W.

As a consequence, for our particular choice of exponential weight function (21), the radially symmetric bump is monotonically decreasing in r, as it will be recover in our numerical experiments in section 6.

5.2 Proof of theorem 5.1.1

The proof of theorem 5.1.1 goes in four steps. First we introduce some notations and recall some basic properties of the Fourier transform in the Poincaré disk. Second we prove two propositions. Third we state a technical lemma on hypergeometric functions, the proof being given in lemma F.0.4 of the appendix F. The last step is devoted to the conclusion of the proof.

5.2.1 First step

In order to calculate Inline graphic, we use the Fourier transform in Inline graphic which has already been introduced in section 4. First we rewrite Inline graphic as a convolution product:

Proposition 5.2.1. For all Inline graphic:

graphic file with name 2190-8567-1-4-i260.gif (26)

Proof. We start with the definition of Inline graphic and use the convolutional form of the integral:

graphic file with name 2190-8567-1-4-i261.gif

In [22], Helgason proves an inversion formula for the hyperbolic Fourier transform and we apply this result to W:

graphic file with name 2190-8567-1-4-i262.gif

the last equality is a direct application of lemma 4.2.1 and we can deduce that

graphic file with name 2190-8567-1-4-i263.gif (27)

Finally we have:

graphic file with name 2190-8567-1-4-i264.gif

which is the desired formula.   □

It appears that the study of Inline graphic consists in calculating the convolution product Inline graphic.

Proposition 5.2.2. For all z = k ·O for k G = SU(1, 1) we have:

graphic file with name 2190-8567-1-4-i266.gif

Proof. Let z = k ·O for k G we have:

graphic file with name 2190-8567-1-4-i267.gif

for all g, k G, Inline graphicλ(g-1k ·O) = Inline graphicλ(k-1g ·O) so that:

graphic file with name 2190-8567-1-4-i268.gif

5.2.2 Second step

In this part, we prove two results:

• the mapping Inline graphic is a radial function, i.e. it depends only upon the variable r.

• the following equality holds for z = tanh(r)e:

graphic file with name 2190-8567-1-4-i270.gif

Proposition 5.2.3. If z = k ·O and z is written Inline graphic with r = d2(z, O) in hyperbolic polar coordinates the function Inline graphic depends only upon the variable r.

Proof. If Inline graphic, then z = rotθ ar ·O and k-1 = a-rrot-θ. Similarly z' = rotθ' ar'·O. We can write thanks to the previous proposition 5.2.2:

graphic file with name 2190-8567-1-4-i274.gif

which, as announced, is only a function of r.   □

We now give an explicit formula for the integral Inline graphic.

Proposition 5.2.4. For all Inline graphic we have:

graphic file with name 2190-8567-1-4-i277.gif

Proof. We first recall a formula from [22].

Lemma 5.2.1. For all g G the following equation holds:

graphic file with name 2190-8567-1-4-i278.gif

Proof. See [22].   □

It follows immediately that for all Inline graphic and r ∈ ℝ we have:

graphic file with name 2190-8567-1-4-i279.gif

We integrate this formula over the hyperbolic ball Bh(0, ω) which gives:

graphic file with name 2190-8567-1-4-i280.gif

and we exchange the order of integration:

graphic file with name 2190-8567-1-4-i281.gif

We note that the integral Inline graphic does not depend upon the variable b = e. Indeed:

graphic file with name 2190-8567-1-4-i283.gif

and indeed the integral does not depend upon the variable b:

graphic file with name 2190-8567-1-4-i284.gif

Finally, we can write:

graphic file with name 2190-8567-1-4-i285.gif

because Φλ = Φ(as solutions of the same equation).

This completes the proof that:

graphic file with name 2190-8567-1-4-i286.gif

   □

5.2.3 Third step

We state a useful formula.

Lemma 5.2.2. For all ω > 0 the following formula holds:

graphic file with name 2190-8567-1-4-i287.gif

Proof. See lemma F.0.4 of appendix F.   □

5.2.4 The main result

At this point we have proved the following proposition thanks to propositions 5.2.1 and 5.2.4.

Proposition 5.2.5. If Inline graphic, Inline graphicis given by the following formula:

graphic file with name 2190-8567-1-4-i289.gif

where

graphic file with name 2190-8567-1-4-i290.gif

We are now in a position to obtain the analytic form for Inline graphic of theorem 5.1.1. We prove that

graphic file with name 2190-8567-1-4-i291.gif

Indeed, in hyperbolic polar coordinates, we have:

graphic file with name 2190-8567-1-4-i292.gif

On the other hand:

graphic file with name 2190-8567-1-4-i293.gif

This yields

graphic file with name 2190-8567-1-4-i294.gif

and we use lemma (5.2.2) to establish (24).

5.3 Linear stability analysis

We now analyse the evolution of small time-dependent perturbations of the hyperbolic stationary-pulse solution through linear stability analysis. We use classical tools already developped in [31,29].

5.3.1 Spectral analysis of the linearized operator

Equation (20) is linearized about the stationary solution V(r) by introducing the time-dependent perturbation:

graphic file with name 2190-8567-1-4-i295.gif

This leads to the linear equation:

graphic file with name 2190-8567-1-4-i296.gif

We separate variables by setting ϕ(z, t) = ϕ(z)eβt to obtain the equation:

graphic file with name 2190-8567-1-4-i297.gif

Introducing the hyperbolic polar coordinates Inline graphic and using the result:

graphic file with name 2190-8567-1-4-i299.gif

we obtain:

graphic file with name 2190-8567-1-4-i300.gif

Note that we have formally differentiated the Heaviside function, which is permissible since it arises inside a convolution. One could also develop the linear stability analysis by considering perturbations of the threshold crossing points along the lines of Amari [20]. Since we are linearizing about a stationary rather than a traveling pulse, we can analyze the spectrum of the linear operator without the recourse to Evans functions.

With a slight abuse of notation we are led to study the solutions of the integral equation:

graphic file with name 2190-8567-1-4-i301.gif (28)

where the following equality derives from the definition of the hyperbolic distance in equation (25):

graphic file with name 2190-8567-1-4-i302.gif

Essential spectrum If the function ϕ satisfies the condition

graphic file with name 2190-8567-1-4-i303.gif

then equation (28) reduces to:

graphic file with name 2190-8567-1-4-i304.gif

yielding the eigenvalue:

graphic file with name 2190-8567-1-4-i305.gif

This part of the essential spectrum is negative and does not cause instability.

Discrete spectrum If we are not in the previous case we have to study the solutions of the integral equation (28).

This equation shows that ϕ(r, θ) is completely determined by its values ϕ(ω, θ) on the circle of equation r = ω. Hence, we need only to consider r = ω, yielding the integral equation:

graphic file with name 2190-8567-1-4-i306.gif

The solutions of this equation are exponential functions eγθ, where γ satisfies:

graphic file with name 2190-8567-1-4-i307.gif

By the requirement that ϕ is 2π-periodic in θ, it follows that γ = in, where n ∈ ℤ. Thus the integral operator with kernel Inline graphic has a discrete spectrum given by:

graphic file with name 2190-8567-1-4-i309.gif

βn is real since:

graphic file with name 2190-8567-1-4-i310.gif

Hence,

graphic file with name 2190-8567-1-4-i311.gif

We can state the following proposition:

Proposition 5.3.1. Provided that for all n ≥ 0, βn < 0 then the hyperbolic stationary pulse is stable.

We now derive a reduced condition linking the parameters for the stability of hyperbolic stationary pulse.

Reduced condition Since Inline graphic is a positive function of r, it follows that:

Stability of the hyperbolic stationary pulse requires that for all n ≥ 0, βn < 0. This can be rewritten as:

graphic file with name 2190-8567-1-4-i313.gif

Using the fact that βn β0 for all n ≥ 1, we obtain the reduced stability condition:

graphic file with name 2190-8567-1-4-i314.gif

Where

graphic file with name 2190-8567-1-4-i315.gif

From (22) we have:

graphic file with name 2190-8567-1-4-i316.gif

Where

graphic file with name 2190-8567-1-4-i317.gif

We have previously established that Inline graphic and I'(ω) is negative by definition. Hence, letting Inline graphic we have

graphic file with name 2190-8567-1-4-i320.gif

By substitution we obtain another form of the reduced stability condition:

graphic file with name 2190-8567-1-4-i321.gif (29)

We also have:

graphic file with name 2190-8567-1-4-i322.gif

and

graphic file with name 2190-8567-1-4-i323.gif

showing that the stability condition (29) is satisfied when N'(ω) > 0 and is not satisfied when N'(ω) > 0.

Proposition 5.3.2 (Reduced condition). If N'(ω) > 0 then for all n ≥ 0, βn < 0 and the hyperbolic stationary pulse is stable.

6 Numerical results

The aim of this section is to numerically solve (13) for different values of the parameters. This implies developing a numerical scheme that approaches the solution of our equation, and proving that this scheme effectively converges to the solution.

Since equation (13) is defined on Inline graphic, computing the solutions on the whole hyperbolic disk has the same complexity as computing the solutions of usual Euclidean neural field equations defined on ℝ2. As most authors in the Euclidean case [31,27,26,29], we reduce the domain of integration to a compact region of the hyperbolic disk. Practically, we work in the Euclidean ball of radius a = 0.5 and center 0. Note that a Euclidean ball centered at the origin is also a centered hyperbolic ball, their radii being different.

We have divided this section into four parts. The first part is dedicated to the study of the discretization scheme of equation (13). In the following two parts, we study the solutions for different connectivity functions: an exponential function, section 6.2, and a difference of Gaussians, section 6.3.

6.1 Numerical schemes

Let us consider the modified equation of (13):

graphic file with name 2190-8567-1-4-i325.gif (30)

We assume that the connectivity function satisfies the conditions (C1)-(C2). Moreover we express z in (Euclidean) polar coordinates such that Inline graphic and Inline graphic. The integral in equation (30) is then:

graphic file with name 2190-8567-1-4-i328.gif

We define Inline graphic to be the rectangle Inline graphic.

6.1.1 Discretization scheme

We discretize Inline graphic in order to turn (30) into a finite number of equations. For this purpose we introduce Inline graphic and Inline graphic,

graphic file with name 2190-8567-1-4-i333.gif

and obtain the (N + 1) (M +1) equations:

graphic file with name 2190-8567-1-4-i334.gif

which define the discretization of (30):

graphic file with name 2190-8567-1-4-i335.gif (31)

where Inline graphic. Similar definitions apply Inline graphic and Inline graphic. Moreover:

graphic file with name 2190-8567-1-4-i339.gif

Inline graphic is the space of the matrices of size n × p with real coefficients. It remains to discretize the integral term. For this as in [33], we use the rectangular rule for the quadrature so that for all Inline graphic we have:

graphic file with name 2190-8567-1-4-i342.gif

We end up with the following numerical scheme, where Inline graphic (resp. Inline graphic) is an approximation of Inline graphic (resp. Inline graphic), Inline graphic:

graphic file with name 2190-8567-1-4-i348.gif

With Inline graphic

6.1.2 Discussion

We discuss the error induced by the rectangular rule for the quadrature. Let f be a function which is Inline graphic on a rectangular domain [a, b] × [c, d]. If we denote by Ef this error, then Inline graphic where m and n are the number of subintervals used and Inline graphic where, as usual, α is a multi-index. As a consequence, if we want to control the error, we have to impose that the solution is, at least, Inline graphic in space.

Four our numerical experiments we use the specific function ode45 of Matlab which is based on an explicit Runge-Kutta (4,5) formula (see [34] for more details on Runge-Kutta methods).

We can also establish a proof of the convergence of the numerical scheme which is exactly the same as in [33] excepted that we use the theorem of continuous dependence of the solution for ordinary differential equations.

6.2 Purely excitatory exponential connectivity function

In this subsection, we give some numerical solutions of (13) in the case where the connectivity function is an exponential function, Inline graphic, with b a positive parameter. Only excitation is present in this case. In all the experiments we set α = 0.1 and Inline graphic with μ = 10.

Constant input We fix the external input I(z) to be of the form:

graphic file with name 2190-8567-1-4-i356.gif

In all experiments we set Inline graphic and σ = 0.05, this means that the input has a sharp profile centered at 0.

We show in Figure 2 plots of the solution at time T = 2500 for three different values of the width b of the exponential function. When b = 1, the whole network is highly excited, whereas as b changes from 1 to 0.1 the amplitude of the solution decreases, and the area of high excitation becomes concentrated around the external input.

Figure 2.

Figure 2

Plots of the solution of equation (13) at T = 2500 for the values μ = 10, α = 0.1 and for decreasing values of the width b of the connectivity, see text.

Variable input In this paragraph, we allow the external current to depend upon the time variable. We have:

graphic file with name 2190-8567-1-4-i358.gif

where Inline graphic. This is a bump rotating with angular velocity Ω0 around the circle of radius r0 centered at the origin. In our numerical experiments we set r0 = 0.4, Ω0 = 0.01, Inline graphic and σ = 0.05. We plot in Figure 3 the solution at different times T = 100, 150, 200, 250.

Figure 3.

Figure 3

Plots of the solution of equation (13) in the case of an exponential connectivity function with b = 0.1 at different times with a time-dependent input, see text.

High gain limit We consider the high gain limit μ → ∞ of the sigmoid function and we propose to illustrate section 5 with a numerical simulation. We set α = 1, κ = 0.04, ω = 0.18. We fix the input to be of the form:

graphic file with name 2190-8567-1-4-i360.gif

with Inline graphic and σ = 0.05. Then the condition of existence of a stationary pulse (23) is satisfied, see Figure 1. We plot a bump solution according to (23) in Figure 4.

Figure 4.

Figure 4

Plot of a bump solution of equation (22) for the values α = 1, κ = 0.04, ω = 0.18 and for b = 0.2 for the width of the connectivity, see text.

6.3 Excitatory and inhibitory connectivity function

We give some numerical solutions of (13) in the case where the connectivity function is a difference of Gaussians, which features an excitatory center and an inhibitory surround:

graphic file with name 2190-8567-1-4-i362.gif

We illustrate the behaviour of the solutions when increasing the slope μ of the sigmoid. We set the Inline graphic so that it is equal to 0 at the origin and we choose the external input equal to zero, I(z, t) = 0. In this case the constant function equal to 0 is a solution of (13).

For small values of the slope μ, the dynamics of the solution is trivial: every solution asymptotically converges to the null solution, as shown in top left hand corner of Figure 5 with μ = 1. When increasing μ, the stability bound, found in subsection 4.5 is no longer satisfied and the null solution may no longer be stable. In effect this solution may bifurcate to other, more interesting solutions. We plot in Figure 5, some solutions at T = 2500 for different values of μ (μ = 3, 5, 10, 20 and 30). We can see exotic patterns which feature some interesting symmetries. The formal study of these bifurcated solutions is left for future work.

Figure 5.

Figure 5

Plots of the solutions of equation (13) in the case where the connectivity function is the difference of two Gaussians at time T = 2500 for α = 0.1 and for increasing values of the slope μ of the sigmoid, see text.

7 Conclusion

In this paper, we have studied the existence and uniqueness of a solution of the evolution equation for a smooth neural mass model called the structure tensor model. This model is an approach to the representation and processing of textures and edges in the visual area V1 which contains as a special case the well-known ring model of orientations (see [1,2,19]). We have also given a rigorous functional framework for the study and computation of the stationary solutions to this nonlinear integro-differential equation. This work sets the basis for further studies beyond the spatially periodic case studied in [15], where the hypothesis of spatial periodicity allows one to replace the unbounded (hyperbolic) domain by a compact one, hence making the functional analysis much simpler.

We have completed our study by constructing and analyzing spatially localised bumps in the high-gain limit of the sigmoid function. It is true that networks with Heaviside nonlinearities are not very realistic from the neurobiological perspective and lead to difficult mathematical considerations. However, taking the high-gain limit is instructive since it allows the explicit construction of stationary solutions which is impossible with sigmoidal nonlinearities. We have constructed what we called a hyperbolic radially symmetric stationary-pulse and presented a linear stability analysis adapted from [31]. The study of stationary solutions is very important as it conveys information for models of V1 that is likely to be biologically relevant. Moreover our study has to be thought of as the analog in the case of the structure tensor model to the analysis of tuning curves of the ring model of orientations (see [1,2,19,35]). However, these solutions may be destabilized by adding lateral spatial connections in a spatially organized network of structure tensor models; this remains an area of future investigation. As far as we know, only Bressloff and coworkers looked at this problem (see [3,11-14,4]).

Finally, we illustrated our theoretical results with numerical simulations based on rigorously defined numerical schemes. We hope that our numerical experiments will lead to new and exciting investigations such as a thorough study of the bifurcations of the solutions of our equations with respect to such parameters as the slope of the sigmoid and the width of the connectivity function.

Competing interests

The authors declare that they have no competing interests.

A Isometries of Inline graphic

We briefly descrbies the isometries of Inline graphic, i.e the transformations that preserve the distance d2. We refer to the classical textbooks in hyperbolic goemetry for details, e.g, [17]. The direct isometries (preserving the orientation) in Inline graphic are the elements of the special unitary group, noted SU(1, 1), of 2 × 2 Hermitian matrices with determinant equal to 1. Given:

graphic file with name 2190-8567-1-4-i364.gif

an element of SU(1, 1), the corresponding isometry γ in Inline graphic is defined by:

graphic file with name 2190-8567-1-4-i365.gif (32)

Orientation reversing isometries of Inline graphic are obtained by composing any transformation (32) with the reflection Inline graphic. The full symmetry group of the Poincaré disc is therefore:

graphic file with name 2190-8567-1-4-i367.gif

Let us now describe the different kinds of direct isometries acting in Inline graphic. We first define the following one parameter subgroups of SU(1, 1):

graphic file with name 2190-8567-1-4-i368.gif

Note that Inline graphic and also ar·O = tanh r, with O being the center of the Poincaré disk that is the point represented by z = 0.

The group K is the orthogonal group O(2). Its orbits are concentric circles. It is possible to express each point Inline graphic in hyperbolic polar coordinates: Inline graphic and r = d2(z, 0).

The orbits of A converge to the same limit points of the unit circle Inline graphic, b±1 = ±1 when r → ±∞. They are circular arcs in Inline graphic going through the points b1 and b-1.

The orbits of N are the circles inside Inline graphic and tangent to the unit circle at b1. These circles are called horocycles with base point b1. N is called the horocyclic group. It is also possible to express each point Inline graphic in horocyclic coordinates: zsar·O, where ns are the transformations associated with the group N (s ∈ ℝ) and ar the transformations associated with the subroup A (r ∈ ℝ).

Iwasawa decomposition The following decomposition holds, see [36]:

graphic file with name 2190-8567-1-4-i373.gif

This theorem allows us to decompose any isometry of Inline graphic as the product of at most three elements in the groups, K, A and N.

B Volume element in structure tensor space

Let Inline graphic be a structure tensor

graphic file with name 2190-8567-1-4-i374.gif

Δ2 its determinant, Δ ≥ 0. Inline graphic can be written

graphic file with name 2190-8567-1-4-i375.gif

Where Inline graphic has determinant 1. Let z = z1 + iz2 be the complex number representation of Inline graphic in the Poincaré disk Inline graphic. In this part of the appendix, we present a simple form for the volume element in full structure tensor space, when parametrized as (Δ, z).

Proposition B.0.1. The volume element in (Δ, z1, z2) coordinates is

graphic file with name 2190-8567-1-4-i377.gif (33)

Proof. In order to compute the volume element in (Δ, z1, z2) space, we need to express the metric Inline graphic in these coordinates. This is obtained from the inner product in the tangent space Inline graphic at point Inline graphic of SDP(2). The tangent space is the set S(2) of symmetric matrices and the inner product is defined by:

graphic file with name 2190-8567-1-4-i380.gif

We note that Inline graphic. We write g instead of Inline graphic. A basis of Inline graphic (or Inline graphic for that matter) is given by:

graphic file with name 2190-8567-1-4-i384.gif

and the metric is given by:

graphic file with name 2190-8567-1-4-i385.gif

The determinant Inline graphic of Inline graphic is equal to G6, where G is the determinant of Inline graphic. G is found to be equal to 2. The volume element is thus:

graphic file with name 2190-8567-1-4-i388.gif

We then use the relations:

graphic file with name 2190-8567-1-4-i389.gif

where Inline graphic, i = 1, 2, 3 is given by:

graphic file with name 2190-8567-1-4-i391.gif

The determinant of the Jacobian of the transformation (x1, x2, x3) → (Δ, z1, z2) is found to be equal to:

graphic file with name 2190-8567-1-4-i392.gif

Hence, the volume element in (Δ, z1, z2) coordinates is

graphic file with name 2190-8567-1-4-i393.gif

   □

C Global existence of solutions

Theorem C.0.1. Let Inline graphic be an open connected set of a real Banach space Inline graphic and J be an open interval of ℝ. We consider the initial value problem:

graphic file with name 2190-8567-1-4-i396.gif (34)

We suppose that Inline graphic and is locally Lipschitz with respect to its second argument. Then for all Inline graphic, there exists τ > 0 and Inline graphic unique solution of (34).

Lemma C.0.1. Under hypotheses of theorem C.0.1, if Inline graphic and Inline graphic are two solutions and if there exists t0 J1 J2 such that V1(t0) = V2(t0) then:

graphic file with name 2190-8567-1-4-i402.gif

This lemma shows the existence of a larger interval J0 on which the initial value problem (34) has a unique solution. This solution is called the maximal solution.

Theorem C.0.2. Under hypotheses of theorem C.0.1, let Inline graphic be a maximal solution. We denote by b the upper bound of J and β the upper bound of J0. Then either β = b or for all compact set Inline graphic, there exists η < β such that:

graphic file with name 2190-8567-1-4-i405.gif

We have the same result with the lower bounds.

Theorem C.0.3. We suppose Inline graphic and is globally Lipschitz with respect to its second argument. Then for all Inline graphic, there exists a unique Inline graphic solution of (34).

D Proof of lemma 3.1.1

Lemma D.0.2. When W is only a function of Inline graphic, then Inline graphic does not depend upon the variable Inline graphic.

Proof. We work in (z, Δ) coordinates and we begin by rewriting the double integral (6) for all Inline graphic:

graphic file with name 2190-8567-1-4-i412.gif

The change of variable Inline graphic yields:

graphic file with name 2190-8567-1-4-i414.gif

And it establishes that Inline graphic does not depend upon the variable Δ. To finish the proof, we show that the following integral does not depend upon the variable Inline graphic:

graphic file with name 2190-8567-1-4-i416.gif (35)

where f is a real-valued function such that Ξ(z) is well defined.

We express z in horocyclic coordinates: zsar.O (see appendix A) and (35) becomes:

graphic file with name 2190-8567-1-4-i417.gif

With the change of variable s s' = −xe2r, this becomes:

graphic file with name 2190-8567-1-4-i418.gif

The relation Inline graphic (proved e.g. in [22]) yields:

graphic file with name 2190-8567-1-4-i420.gif

with Inline graphic and Inline graphic, which shows that Ξ(z) does not depend upon the variable z, as announced.   □

E Proof of lemma 3.1.2

In this section we prove the following lemma.

Lemma E.0.3. When W is the following Mexican hat function:

graphic file with name 2190-8567-1-4-i423.gif

where:

graphic file with name 2190-8567-1-4-i424.gif

with 0 ≤ σ1 σ2 and 0 ≤ A ≤ 1.

Then:

graphic file with name 2190-8567-1-4-i425.gif

where erf is the error function defined as:

graphic file with name 2190-8567-1-4-i426.gif

Proof. We consider the following double integrals:

graphic file with name 2190-8567-1-4-i427.gif (36)

so that:

graphic file with name 2190-8567-1-4-i428.gif

Since the variables are separable, we have:

graphic file with name 2190-8567-1-4-i429.gif

One can easily see that:

graphic file with name 2190-8567-1-4-i430.gif

We now give a simplified expression for Ξi. We set Inline graphic and then we have, because of lemma 3.1.1:

graphic file with name 2190-8567-1-4-i432.gif

The change of variable x = arctanh(r) implies Inline graphic and yields:

graphic file with name 2190-8567-1-4-i434.gif

then we have a simplified expression for Ξi:

graphic file with name 2190-8567-1-4-i435.gif

   □

F Proof of lemma 5.2.2

Lemma F.0.4. For all ω > 0 the following formula holds:

graphic file with name 2190-8567-1-4-i436.gif

Proof. We write z in hyperbolic polar coordinates, z = tanh (r) e(see appendix A). We have:

graphic file with name 2190-8567-1-4-i437.gif

Because of the above definition of Φλ, this reduces to

graphic file with name 2190-8567-1-4-i438.gif

In [22] Helgason proved that:

graphic file with name 2190-8567-1-4-i439.gif

with Inline graphic. We then use the formula obtained by Erdelyi in [32]:

graphic file with name 2190-8567-1-4-i441.gif

Using some simple hyperbolic trigonometry formulae we obtain:

graphic file with name 2190-8567-1-4-i442.gif

from which we deduce

graphic file with name 2190-8567-1-4-i443.gif

Finally we use the equality shown in [32]:

graphic file with name 2190-8567-1-4-i444.gif

In our case we have: a = ν, b = 1 - ν, c = 2 and z = - sinh (ω)2, so Inline graphic. We obtain

graphic file with name 2190-8567-1-4-i446.gif

Since Hypergometric functions are symmetric with respect to the first two variables:

graphic file with name 2190-8567-1-4-i447.gif

we write

graphic file with name 2190-8567-1-4-i448.gif

which yields the announced formula

graphic file with name 2190-8567-1-4-i449.gif

Contributor Information

Gregory Faye, Email: gregory.faye@inria.fr.

Pascal Chossat, Email: gregory.faye@inria.fr.

Olivier Faugeras, Email: gregory.faye@inria.fr.

Acknowledgements

This work was partially funded by the ERC advanced grant NerVi number 227747.

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