Attractor and saddle node dynamics in heterogeneous neural fields
 Peter beim Graben^{1, 2} and
 Axel Hutt^{3}Email author
DOI: 10.1140/epjnbp17
© beim Graben and Hutt; licensee Springer. 2014
Received: 21 November 2013
Accepted: 26 March 2014
Published: 9 May 2014
Abstract
Background
We present analytical and numerical studies on the linear stability of spatially nonconstant stationary states in heterogeneous neural fields for specific synaptic interaction kernels.
Methods
The work shows the linear stabiliy analysis of stationary states and the implementation of a nonlinear heteroclinic orbit.
Results
We find that the stationary state obeys the Hammerstein equation and that the neural field dynamics may obey a saddlenode bifurcation. Moreover our work takes up this finding and shows how to construct heteroclinic orbits built on a sequence of saddle nodes on multiple hierarchical levels on the basis of a LotkaVolterra population dynamics.
Conclusions
The work represents the basis for future implementation of metastable attractor dynamics observed experimentally in neural population activity, such as Local Field Potentials and EEG.
Keywords
Chaotic itinerancy Linear stability Heteroclinic orbits LotkaVolterra modelBackground
Neural field models, such as [1, 2] are continuum limits of largescale neural networks. Typically, their dynamic variables describe either mean voltage [2] or mean firing rate [1, 3] of a population element of neural tissue (see [4, 5] for recent reviews).
where K(x, y) is the spatial synaptic connectivity between site y ∈ Ω and site x ∈ Ω, and S is a nonlinear, typically sigmoidal, transfer function. This model neglects external inputs for simplicity but without constraining the generality of the subsequent results. Possible synaptic time scales are supposed to be included in the kernel function K and can be introduced by a simple scaling of time.
In general, the connectivity kernel K(x, y) fully depends on both sites x and y, which case is referred to as spatial heterogeneity. If the connectivity solely depends on the difference between x and y, i.e. K(x, y) = K(x  y), the kernel is called spatially homogeneous[2]. Furthermore, if the connectivity depends on the distance between x and y only, i.e. K(x, y) = K(x  y), with x as some norm in Ω, the kernel is spatially homogeneous and isotropic[6].
Spatially homogeneous (respectively isotropic) kernels have been intensively studied in the literature due to their nice analytical properties. In this case, the evolution equations have exact solutions such as bumps [2, 7], breathers [8–10] or traveling waves [7, 11]. Moreover, such kernels allow the application of the technique of Green’s functions for deriving partial neural wave equations [7, 12, 13].
The present work focuses on spatially heterogeneous neural fields which have been discussed to a much lesser extent in previous studies than homogeneous neural fields [14–22]. This study resumes these attempts by investigating stationary states of the Amari equation (1) with heterogeneous kernels and their stability. Such a theory would be mandatory for modeling transient neurodynamics as is characteristic, e.g., for human cognitive phenomena [23], human early evoked potentials [24] or, among many other phenomena, for bird songs [25].
The article is structured in the following way. In the “Results” section we present new analytical results on stationary solutions of the Amari equation (1) and their stability in the presence of heterogeneous connectivity kernels. Moreover, we present numerical simulation results for the kernel construction and its stability analysis. The “Methods” section is devoted to construct such kernels through dyadic products of desired stationary states (cf. the previous work of Veltz and Faugeras [26]). A subsequent linear stability analysis reveals that these stationary solutions could be either attractors or saddles, depending on the chosen parametrization. Finally, we present a way to connect such saddle state solutions via heteroclinic sequences [27, 28] in order to construct transient processes.
Results
In this section we present the main results of our study on heterogeneous neural fields.
Stationary states and their stability
Analytical study
which indicates immediately a method to find a nontrivial solution numerically as shown below.
where $L(x,y)=K(x,y){S}^{\prime}\left(y\right)$ and ${S}^{\prime}\left(y\right)=\mathrm{d}S\left[\phantom{\rule{0.3em}{0ex}}{V}_{0}\right(y\left)\right]/\mathrm{d}{V}_{0}\left(y\right)$. A linear stability analysis of (6) carried out in the next section shows that a nontrivial stationary state V_{0}(x) is either a fixed point attractor, or (neglecting a singular case) a saddle with onedimensional unstable manifold. Such saddles can be connected to form stable heteroclinic sequences.
Numerical study
To gain deeper insight into possible stationary solutions of the Amari equation (1) and their stability, the subsequent section presents the numerical integration of equation (1) in one spatial dimension for a specific spatial synaptic connectivity kernel.
parameterized by the amplitude W_{0}, the variance σ^{2}, the noise level κ and the spatial discretization interval Δ x.
For each stationary state, one obtains a kernel L(x, y) of the linear stability analysis whose spectrum characterizes the stability of the system in the vicinity of the stationary state. If the eigenvalue with the maximum real part ε_{1} > 1, then the stationary state V_{0}(x) is exponentially unstable whereas ε_{1} < 1 guarantees exponential stability. Figure 1(b) shows the eigenmode e_{1}(x) corresponding to the eigenvalue with maximum real part which has a similar shape as the stationary state.
Taking a closer look at the stability of V_{0}(x), the computation of the eigenvalues ε_{ k }, k = 1, …, n reveals a dramatic gap in the spectrum: the eigenvalue with maximum real part ε_{1} is well isolated from the rest of the spectrum $\left\{{\epsilon}_{k>1}\right\}$ with ε_{k>1} < 10^{14}. This is in accordance to the discussion of Eq. (17) on the linear spectrum.
Recalling the presence of the trivial stable solution V = 0, the activity shown in Figure 4 indicates the bistability of the system for the given parameter set.
Heteroclinic orbits
The previous section has shown that heterogeneous neural fields may exhibit various stationary states with different stability properties. In particular we found that stationary states could be saddles with onedimensional unstable manifolds that could be connected to stable heteroclinic sequences (SHS: [27, 28]) which is supported by experimental evidence [24, 32, 33]. We present in the following paragraphs our main findings on heterogeneous neural fields exhibiting heteroclinic sequences and also hierarchies of such sequences.
One level heteroclinic sequence
The solution of Eq. (8) represents a heteroclinic sequence that connects saddle points {V_{ k }(x)} along their respective stable and unstable manifolds. Its transient evolution is described as winnerless competition in a LotkaVolterra population dynamics governed by interaction weights ρ_{ i k } between neural populations k and i and their respective growth rates σ_{ i }.
In Eq. (9) the $\left\{{V}_{k}^{+}\right(x\left)\right\}$ comprise a biorthogonal system of the saddles {V_{ k }(x)}. Therefore, the kernel K_{1}(x, y) describes a Hebbian synapse between sites y and x that has been trained with pattern sequence V_{ k }. This finding confirms the previous result of [18]. Moreover the threepoint kernel K_{2}(x, y, z) further generalizes Hebbian learning to interactions between three sites x, y, z ∈ Ω. Note that the kernels K_{ i } are linear combinations of dyadic product kernels, similar to those introduced in (3). Thus, our construction of heteroclinic orbits straightforwardly results in PincherleGoursat kernels used in [26].
Multilevel hierarchy of heteroclinic sequences
with a tensor product kernel $H(s,{s}^{\prime})=(K\otimes G)(s,{s}^{\prime})$ and integration domain $M=\Omega \times ]\infty ,t]$.
Here, the ${V}_{{k}_{\nu}}^{\left(\nu \right)}\left(x\right)$ denote the k_{ ν }th saddle in the νth level of the hierarchy (containing n_{ ν } stationary states). Saddles are chosen again in such a way that they form a system of biorthogonal modes ${V}_{{k}_{\nu}}^{{\left(\nu \right)}^{+}}\left(x\right)$ whose behavior is determined by LotkaVolterra dynamics with growth rates ${\sigma}_{{k}_{\nu}}^{\left(\nu \right)}>0$ and timedependent interaction weights ${\rho}_{{k}_{\nu}{j}_{\nu}}^{\left(\nu \right)}\left(t\right)>0$, ${\rho}_{{k}_{\nu}{k}_{\nu}}^{\left(\nu \right)}\left(t\right)=1$ that are given by linear superpositions of templates ${r}_{{k}_{\nu}{j}_{\nu}{l}_{\nu +1}}^{\left(\nu \right)}$. Additionally, τ^{(ν)} are the characteristic time scales for level ν. Levels are temporally well separated through τ^{(ν)}≫τ^{(ν + 1)}[35].
Interestingly these kernels are timeindependent. Since neural field equations can be written in the same form as Eq. (12), this result shows that hierarchies of LotkaVolterra systems are included in the neural field description. Again we point out that the resulting kernels are linear combinations of dyadic products as introduced in Eq. (3), see also the work of Veltz and Faugeras [26].
Discussion
This study considers spatially heterogeneous neural fields describing the mean potential in neural populations according to the Amari equation [2]. To our best knowledge this work is one of the first deriving the implicit conditions for stationary solutions and identifying the corresponding stability constraint as the Hammerstein integral equation. Moreover, as one of the first studies our work derives conditions for the linear stability of such stationary solutions and derives an analytical expression for stability subjected to the properties of heterogeneous synaptic interaction. The analytical results are complemented by numerical simulations illustrating the stability and instability of heterogeneous stationary states. We point out that the results obtained extend previous studies both on homogeneous neural fields and heterogeneous neural fields as other studies in this context have done before [21].
By virtue of the heterogeneity of the model, it is possible to consider more complex spatiotemporal dynamical behavior than the dynamics close to a single stationary state. We show how to construct hierarchical heteroclinic sequences built of multiple stationary states each exhibiting saddle node dynamics. The work demonstrates in detail how to construct such sequences given the stationary states and their saddle node dynamics involved. Motivated by a previous study on hierarchical heteroclinic sequences [25, 34], we constructed such sequences in heterogeneous neural fields of the Amari type. Our results indicate that such a hierarchy may be present in a single heterogeneous neural field whereas previous studies [25, 34] consider the presence of several neural populations to describe heteroclinic sequences. The kernels obtained from heteroclinic saddle node dynamics are linear superpositions of tensor product kernels, known as PincherleGoursat kernels in the literature [26].
Conclusion
The present work is strongly related to the literature hypothesizing the presence of chaotic itinerant neural activity, cf. previous work by [36, 37]. This concept of chaotic itinerancy is attractive but yet lacking a realistic neural model. We admit that the present work represents just a first initial starting point for further model analysis of sequential neural activity in heterogeneous neural systems. It opens up an avenue of future research and promises to close the gap between the rather abstract concept of sequential, i.e. temporally transient, neural activity and corresponding mathematical neural models.
Methods
Stationary states and their stability
In order to learn more about the spatiotemporal dynamics of a neural field, in general it is reasonable to determine stationary states and to study their linear stability. This already gives some insight into the nonlinear dynamics of the system. Since the dynamics depends strongly on the interaction kernel and the corresponding stationary states, it is necessary to work out conditions for the existence and uniqueness of stationary states and their stability.
Analytical study
and the Nemytskij operator $\mathcal{Fu}\left(x\right)=S\left(u\right(x\left)\right)$. For instance, a simple criterion for the existence of at least a single nontrivial solution is the symmetry and positive definiteness of the kernel K(x, y) and the condition S(u) ≤ C_{1}u + C_{2}[29]. Moreover previous studies have proposed analytical [40] and numerical [41] methods to find solutions of the Hammerstein equation (2).
Since f_{ m } is a nonlinear function of V_{ n }, Eq. (13) has multiple solutions {V_{ n }} for a given biorthogonal basis. Hence, V_{0}(x) may not be unique.
Considering small deviations u(x, t) = V(x, t)  V_{0}(x) from a nontrivial stationary state V_{0}(x), these deviations obey the linear equation (6) with $L(x,y)=K(x,y){S}^{\prime}\left(y\right)$ and ${S}^{\prime}\left(y\right)=\mathrm{d}S\left[\phantom{\rule{0.3em}{0ex}}{V}_{0}\right(y\left)\right]/\mathrm{d}{V}_{0}\left(y\right)$.
From (17) we deduce two important results:

a certain eigenmode ${e}_{{k}_{0}}\left(x\right)$ is proportional to the stationary state V_{0}(x) with scaling factor $\u3008{V}_{0},{e}_{{k}_{0}}{\u3009}_{S}/{\epsilon}_{{k}_{0}}$, and

other eigenmodes in the eigenbasis ${e}_{k\ne {k}_{0}}\left(x\right)$ are orthogonal to V_{0}(x) yielding 〈V_{0},e_{ k }〉_{ S } = ε_{ k } = 0.
Hence for dyadic product kernels (3) the spectrum of L includes one eigenmode with eigenvalue ε_{1} ≠ 0, i.e. λ_{1} ≠ 1, while all other eigenmodes are stable with ε_{k≠1} = 0 and thus λ_{k≠1} = 1. Therefore, a nontrivial stationary state could become either an asymptotically stable fixed point, i.e. an attractor, for ε_{ n } < 1, for all n, or a saddle with a onedimensional unstable manifold, for ε_{1} > 1 and ε_{n≠1} < 1 (neglecting the singular case ε_{1}= 1 ).
inserted into the Hammerstein equation (2) for a dyadic product kernel (3).
Numerical study
We employ an Eulerforward integration scheme for the temporal evolution with discrete time step Δ t = 0.05.
We render the stationary state random by adding the noise term η_{ i } which are random numbers taken from a normal distribution with zero mean and unit variance. We point out that we choose η_{ i } such like $\left\sum _{i=1}^{n}{\eta}_{i}\right<0.05$ in order to not permit an amplitude increase in the dynamics by noise, but just as a modulation. The sigmoid function is chosen as $S\left(V\right)=1/(1+exp(\alpha (V\theta )\left)\right)$ parameterized by the slope parameter α and the mean threshold θ.
Heteroclinic orbits
The general neural field equation (10) supplies the Amari equation (1) as a special case for G(t) = e^{t}Θ(t) (Θ(t) as Heaviside’s step function). Then Q = ∂_{ t } + 1 is the Amari operator for the Green’s function G(t). For secondorder synaptic dynamics [42] and for the filter properties of complex dendritc trees [43], more complicated kernels or differential operators, respectively, have to be taken into account.
with a sequence of integral kernels H_{ m } that can be read off after some further tedious rearrangements [44].
Onelevel hierarchy
with growth rates σ_{ k } > 0, and interaction weights ρ_{ k j } > 0, ρ_{ k k } = 1, that are trained by the algorithm of [27] and [28] for the desired sequence of transitions.
Comparison of the first three terms with (28) yields the result (9).
Multilevel hierarchy of heteroclinic sequences
with growth rates ${\sigma}_{{k}_{\nu}}^{\left(\nu \right)}>0$, and timedependent interaction weights ${\rho}_{{k}_{\nu}{j}_{\nu}}^{\left(\nu \right)}\left(t\right)>0$, ${\rho}_{{k}_{\nu}{k}_{\nu}}^{\left(\nu \right)}\left(t\right)=1$. Again, $\nu \in \mathbb{N}$ indicates the level within the hierarchy, n_{ ν } is the number of stationary states and τ^{(ν)} represents the characteristic time scale of that level. Levels are temporally well separated through τ^{(ν)}≫τ^{(ν + 1)}[35].
where the constants ${r}_{{k}_{\nu}{j}_{\nu}{l}_{\nu +1}}^{\left(\nu \right)}$ serve as parameter templates.
Eventually, we insert all results of the recursion of (36) into the field equation (32) and eliminate the amplitudes ${\alpha}_{{k}_{\nu}}^{\left(\nu \right)}\left(t\right)$ by means of (34) in order to get a series expansion of V(s) in terms of spatiotemporal integrals. This series must equal the generalized Volterra series (21), such that the kernel H(s,s^{′}) can be reconstructed to solve the neural field inverse problem [44]. For more mathematical details on the derivation of the twolevel hierarchy, see Additional file 1.
Declarations
Acknowledgments
This research has been supported by the European Union’s Seventh Framework Programme (FP7/20072013) ERC grant agreement No. 257253 awarded to AH, hosting PbG during fall 2013 in Nancy, and by a Heisenberg fellowship (GR 3711/12) of the German Research Foundation (DFG) awarded to PbG.
Authors’ Affiliations
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