Neural Fields Masses and Bayesian Modelling
Neural Fields Masses and Bayesian Modelling
The Wellcome Trust Centre for Neuroimaging, University College London, Queen Square,
WC1N 3BG
Abstract
This chapter considers the relationship between neural field and mass models
and their application to modelling empirical data. Specifically, we consider neural
masses as a special case of neural fields, when conduction times tend to zero and
focus on two exemplar models of cortical microcircuitry; namely, the Jansen-Rit
and the Canonical Microcircuit Model. Both models incorporate parameters per-
taining to important neurobiological attributes, such as synaptic rate constants and
the extent of lateral connections. We describe these models and show how Bayes-
ian inference can be used to assess the validity of their field and mass variants,
given empirical data. Interestingly, we find greater evidence for neural field vari-
ants in analyses of LFP data but fail to find more evidence for such variants, rela-
tive to their neural mass counterparts, in MEG (virtual electrode) data. The key
distinction between these data is that LFP data are sensitive to a wide range of
spatial frequencies and the temporal fluctuations that these frequencies contain. In
contrast, the lead fields, inherent in non-invasive electromagnetic recordings, are
necessarily broader and suppress temporal dynamics that are expressed in high
spatial frequencies. We present this as an example of how neuronal field and mass
models (hypotheses) can be compared formally.
Introduction
Neural mass models are a particular case of neural fields, where the (hidden
neuronal) states of populations of neurons are functions of time only. Such models
can generate temporal responses from one or several interconnected populations
and have been used successfully to explain empirical electrophysiological data
from local field potentials (LFP) and EEG/MEG (see e.g. Lopes da Silva et al.,
1974;van Rotterdam et al., 1982; Steriade and Deschenes, 1984; Lumer et al.,
1997; Valdes et al.., 1999; Liley et al., 1999; Riera et al.,2007; Moran et al.,
2007; Kiebel et al., 2009). To date, neural mass models have been largely based
upon point sources and formulated using ordinary differential equations (ODEs).
A key challenge in this area has been to model observed signals as being gener-
ated by continuous and spatially distributed neuronal activity, of the sort observed
directly using high density multi-electrode arrays and optical imaging. Here, we
consider how one can address this challenge using neural field models.
Neural field models model current fluxes as continuous processes on a cortical
manifold, using partial differential equations (PDEs) (see Deco et al., 2008 for a
1 http://www.fil.ion.ucl.ac.uk/spm/
3
review and also Nunez, 1995,1996; Nunez et al., 2006; Lopes da Silva and Storm
van Leeuwen, 1978; Attay and Hutt, 2006; Breakspear et al., 2006; Bressloff,
1996,2001;Coombes et al.,2003,2005,2007; Freeman 2003,2005; Liley et al.,
2002; Robinson et al., 2001,2002,2003; O’Connor et al., 2005; Jirsa and Haken,
1996; Qubbaj and Jirsa,2007;Jirsa, 2009; Ghosh et al., 2008). The key advance
that neural field models offer, over conventional neural mass models, is that they
embody spatial parameters (like the density and extent of lateral connections).
This means, in principle, one can estimate the spatial parameters of cortical infra-
structures generating electrophysiological signals (and infer changes in those pa-
rameters over different levels of an experimental factor) from empirical data. This
rests on modelling responses not just in time but also over space. Clearly, to ex-
ploit this sort of model, one needs to measure the temporal dynamics of observed
cortical responses over different spatial scales; for example, with high-density re-
cordings, at the epidural or intracortical level. However, the impact of spatially ex-
tended dynamics is not restricted to expression over space but can also have pro-
found effects on temporal (e.g., spectral) responses at one point (or averaged
locally over the cortical surface). This means that neural field models can also
play a key role in the modelling of non-invasive electrophysiological data that
does not resolve spatial activity directly.
that comprise multiple layers. This allows us to model the spectral activity of cor-
tical fields as measured on the cortical surface and to also compute the corre-
sponding spectral responses, as measured by invasive (LFP) and non-invasive
(MEG) electrophysiological sensors in an efficient manner.
The resulting scheme can be regarded as inverting population models of the
Amari type – using real data – with a particular focus on Bayesian model inver-
sion. Previous work in a similar vein includes the use of Kalman filters to develop
estimation schemes for both neural mass (Valdes et al., 1999; Riera et al., 2007)
and neural field models of a single population (Schiff and Sauer, 2008; Galka et
al., 2008). In a related approach, (Daunizeau et al., 2009) replaced the standard di-
pole source – used in neural mass models – with the principal Fourier mode of a
neural field, for the particular case of exponentially decaying synaptic density over
the cortical surface. Finally, (Markounikau et al., 2010) used a combination of lin-
ear and nonlinear optimisation methods to invert a two-layered neural field model
of voltage-sensitive dye data, describing inhibitory and excitatory populations
(without conduction delays).
Dynamic Causal Modelling (DCM) allows for the comparison and estimation
of biophysically plausible models of fMRI, EEG, MEG and LFP data (Friston et
al., 2003; David et al., 2006; Moran et al., 2007; 2009; 2011). DCM calls on an
underlying generative model to predict observed data. As with other state-space
models, Dynamic Causal Models are based on a combination of evolution equa-
tions for the hidden neuronal states with static observer equations:
V (t ) f (V ,U , )
(1)
Y (t ) L(V , )
V 2 BV B 2V D F V G U
Y (t ) L ( x, ) V ( x, t )dx
v1 (t )
V (t )
vn (t )
In this section, we provide a brief review of the well-known Jansen and Rit
(JR) neural mass model (Jansen and Rit, 1995). In the JR model, each cortical
source is modelled with three subpopulations: excitatory spiny stellate input cells,
inhibitory interneurons and deep excitatory output pyramidal cells (for classical
approaches to modelling such populations with neural fields, see e.g. Freeman
1972; Wilson and Cowan 1973; Amari 1977; Nunez 1995). For simplicity, we
consider a single source, noting that extensions to multiple sources involve adding
extrinsic (between-source) connections or kernels. The JR model is a particular in-
stance of Equation (1) (where the convolution of presynaptic input is second-
order):
A diag me , mi , me
B diag e , i , e
(4)
e me
G 0
0
1
( va ) (6)
1 exp(r ( va ))
Here, r and are parameters that determine the shape of this sigmoid activa-
tion function. In particular, r is synaptic gain and is the postsynaptic potential
at which the half of the maximum firing rate is elicited. In Equation (5),
d ab (vb ) is (endogenous) presynaptic input to the a-th population from the b-th
and corresponds to the mapping F V . See Fig. 1 for a schematic of this model.
Figure 1: This schematic summarizes the equations of motion or state equations that specify a
(Jansen and Rit 1995) neural mass model of a single source. This model contains three populations,
each loosely associated with a specific cortical subpopulation or layer. The second-order differential
equations describe changes in hidden states (e.g., voltage) that subtend observed local field potentials
or EEG signals. These differential equations effectively implement a linear convolution of presynaptic
activity to produce postsynaptic depolarization. Average firing rates within each subpopulation are
then transformed through a nonlinear (sigmoid) voltage-firing rate function () to provide inputs to
other populations. These inputs are weighted by connection strengths.
8
Recent work suggests that superficial layers of visual cortex oscillate preferen-
tially at gamma frequencies, while deep layers primarily oscillate at lower al-
pha/beta frequencies (Buffalo et al., 2011). Since forward connections originate
largely from superficial layers and backward connections primarily originate from
deep layers, these spectral asymmetries suggest that forward connections use
faster (gamma) temporal frequencies, while backward connections may employ
lower (beta) frequencies – a suggestion that has experimental support (Roopun et
al., 2010). These asymmetries mandate something quite remarkable: namely, a
synthesis and segregation of forward and backward output from afferent input.
This segregation can only arise from local neuronal computations that are formally
structured and precisely interconnected. The canonical microcircuit is a detailed
proposal for such a laminar-specific intracortical architecture that describes how
information flows through the cortical column. This model is based on findings in
the primary visual cortex (Douglas and Martin, 1991) but recent work (Lefort et
al., 2009; Weiler et al., 2008) indicates that similar microcircuits exist in other re-
gions – such as somatosensory and motor cortex.
Douglas and Martin recorded intracellular potentials from cells in area 17 of
the cat, while they stimulated cortical afferents, and noticed a strong compartmen-
talization of the superficial and deep cell properties – reflected in slow superficial
responses and fast input layer responses. The authors created a conductance-based
model that reproduced the evolution of excitation and inhibition through the corti-
cal circuit with great precision. This model contained three groups of cells: super-
ficial and deep pyramidal cells, and a common pool of inhibitory cells. All three
pools of neurons receive thalamic drive – although the thalamic drive to deep
layer cells was weaker than the other inputs. All neuronal populations had self-
connectivity, and are interconnected with the other populations. Finally, inhibition
was stronger onto the deep pyramidal population. This model was able to repro-
duce the features observed in their electrophysiological recordings – including the
latency difference between superficial and input layer neurons – and has served to
establish several basic properties that are now believed to be replicated in other
cortical areas: first, although superficial and deep compartments are strongly in-
terconnected, their response properties are also segregated. Second, cortex is not
under tonic inhibition, rather, both excitation and inhibition are generated by af-
ferent thalamic input and both shape ongoing cortical responses. Third, the ca-
nonical microcircuit can amplify thalamic inputs to generate self-sustaining activ-
ity, while also maintaining a delicate balance between excitation and inhibition –
so as to prevent runaway excitation.
Previous computational modelling studies indicate that this circuitry allows the
cortex to optimally organize and integrate bottom-up, lateral, and top-down infor-
mation (Raizada and Grossberg, 2003). Douglas and Martin suggest that the rich
anatomical connectivity of superficial layer 2/3 pyramidal cells allows them to
collect information from all relevant top-down, lateral, and bottom-up inputs, and
– through processing in the dendritic tree – select the most likely interpretation of
its subcortical inputs. For a discussion and more details about the canonical mi-
9
crocircuit and its potential role in predictive coding we refer the reader to (Bastos
et al., under review).
Haeusler and Maass used Hodgkin and Huxley neurons to build a realistic mi-
crocircuit model and showed that a cortical column – whose connectivity con-
forms to the canonical microcircuit – can perform various computations effi-
ciently, in relation to a column with random connectivity (Haeusler and Maass,
2007). By collapsing two pairs of cell types in the Haeusler and Maass model,
while preserving the topology of the connectivity, one obtains the canonical mi-
crocircuit (CMC) depicted in Fig. 2: this circuit comprises four populations: exci-
tatory spiny stellate input cells (1), inhibitory interneurons (2), deep excitatory py-
ramidal cells (3) and superficial excitatory pyramidal cells (4). The corresponding
evolution equations for the neuronal states (the analogues of Equation 5) are (see
also Fig. 2):
Figure 2: This figure shows the evolution equations that specify a Canonical Microcircuit neural
mass model of a single source. This model contains four populations occupying different cortical lay-
ers: the pyramidal cell population of the JR model is here split into two subpopulations allowing a
separation of the sources of forward and backward connections in cortical hierarchies. As with the JR
model, second-order differential equations mediate a linear convolution of presynaptic activity to pro-
10
duce postsynaptic depolarization. This depolarization gives rise to firing rates within each sub-
population that provide inputs to other populations.
Neural Field extensions of the Canonical Microcircuit and Jansen and Rit
models
In this section, we transcribe the neural mass models described above into neu-
ral fields. In the case of fields, we consider spatially extended sources occupying
bounded manifolds (patches) in different layers that lie beneath the cortical sur-
face. In this setting, each subpopulation of a neural mass model now becomes a
layer in the cortical sheet. The dynamics of cortical sources conform to integrodif-
ferential equations, such as the Wilson-Cowan or Amari equations, where cou-
pling is parameterised by matrix-valued coupling kernels – namely, smooth (ana-
lytic) connectivity matrices that also depend on time and space. Assuming that the
connectivity kernels dij ( x, t ) appearing in Equation (2) factorize into
dij ( x, t ) ij (| x |) (t ij | x |) , Equation (2) becomes
where is the inverse speed at which spikes propagate along connections and in-
teractions among populations – within and across macrocolumns – are described
by the connectivity kernel K K ( i ) K ( e ) . This form provides an explicit parame-
terization of conduction delays that will be exploited later, when using the field
model as an observation model. One can see that in the infinite speed limit 0
the spatial convolution in Equation (8) disappears (to within a scaling constant)
and we recover the neural mass Equations (3).
For both the canonical microcircuit and the Jansen and Rit models, the intrinsic
connectivity K ( i ) is an exponentially decaying kernel, commonly used in the lit-
erature to account for excitatory and inhibitory interactions (see e.g. Pinotsis et al.,
2012). The extrinsic connectivity kernel K ( e ) was introduced in (Pinotsis and Fris-
ton, 2011; Grindrod and Pinotsis, 2011) to model patchy lateral (horizontal) con-
nections and is characterized by non-central peaks allowing for differences in (and
estimation of) the range and dispersion of lateral connections, summarized in
terms of the parameters ha and caa respectively. The canonical microcircuit
(CMC) field model we consider here is described by Equation (8), where the con-
nectivity kernel is given by
11
K K (i ) K ( e )
k11( i ) k12(i ) 0 k14( i ) k11( e ) 0 0 0
(i )
k k22(i ) k23(i ) 0 0 ( e)
k 0 0
K (i ) 21 , K (e) 22
0 k32(i ) k33(i ) 0 0 0 k33( e ) 0
(i )
k41 0 k44
(i) (e)
0 0 k44 0 0
(i )
kab ( x) 12 aab e cab | x|
kaa( e ) ( x) 12 caa e caa x ha
e
caa x ha
(9)
Here, the parameters aab and cab encode the strength (analogous to the num-
ber
(9) of synaptic connections) and extent of intrinsic connections between cortical
layers. The intrinsic connections can be regarded as inter-laminar connections
within a macrocolumn, while the extrinsic – between macrocolumn connections –
correspond to horizontal connections and connect layers of the same type at a dis-
tance ha , see Fig. 3.
Figure 3: Connectivity kernel describing a combination of patchy but isotropic distributions by us-
ing connectivity kernels with non-central peaks. This kernel models sparse intrinsic connections in cor-
tical circuits that mediate both local and non-local interactions. In other words, neurons talk both to
their immediate neighbours and receive input from remote populations who share the same functional
selectivity; see Equation (9). The insert is a modified from www.ini.uzh.ch/node/23776
12
This kernel models sparse intrinsic connections in cortical circuits that mediate
both local and non-local interactions and allows one to estimate properties of lat-
eral interactions that are particularly relevant in the context of data obtained from
retinotopically mapped visual cortex. We will see an example of this using an at-
tention task and MEG data below. On the other hand, to illustrate the Jansen and
Rit field model in non-mapped cortex, we will use data from the auditory cortex
under anaesthesia and neglect extrinsic connectivity; that is K ( e ) 0 . In this ex-
ample, we will use the same exponential decaying form, as for the CMC field
model above, namely the function K ( x) K ( i ) ( x) where
0 0 k13(i )
K (i ) 0 0 k 23(i ) (10)
(i ) (i )
k 31 k 32 0
The predicted response at an LFP or virtual electrode – for a given set of neural
and lead field parameters is obtained by integrating over the cortical patch
g ( , ) Y ( , )Y * ( , )
| L (k , ) |2 QT (k , ) g u (k , )T (k , )* Q T
(14)
k
2
where g u (k , ) U (k , ) is the auto-power spectrum of external input. The
above expression depends upon the transfer function T (k , ) associated with the
evolution Equations of the model, which we will derive below.
For the case of MEG virtual electrodes and LFP data we consider here, the gain
function has a simple Gaussian form, which we parameterize in terms of its dis-
persion such that L( x, ) e x / 2 – noting that the amplitude is fixed to avoid
2 2
redundancy with the parameters Q . This leads to Fourier coefficients of the form
L(k , ) e 2 k
2 2 2
and Equation (14) becomes
g ( , ) e 2
2
k
QT (k , ) g u (k , )T (k , )* QT
2 2 2
(15)
k
The function T (k , ) can be derived by assuming that the neural field defined by
Equation (2) is perturbed around a spatially homogeneous steady-state V0 at-
tained in the absence of external or exogenous perturbations, (see also Pinotsis and
Friston, 2011; Pinotsis et al., 2012):
Using linear systems analysis, we define the transfer function of a field model
with the following relation
14
P (k , )
T (k , ) (17)
U (k , )
U (k , ) FT (U ( x, t ))
(18)
U ( x, t )e ikx it dtdx
and P(k , ) is the Fourier transform of the perturbations around the steady-state
solution. Given the transfer function, we can characterise the spectral response of
the system to any external input, in terms of the underlying connectivity kernel,
propagation velocities and post-synaptic response function: Substituting
V ( x, t ) V0 P( x, t ) into Equation (2) and expanding F V around V0 , we obtain
a second-order expression for the perturbations P( x, t )
2 BP B 2 P AB D P GU
P
re r
(va 0) ab (19)
ab (1 e r ) 2
vb
0 ab
where is the gain of the nonlinear mapping between depolarisation and firing
rate. Equations (17) and (19) provide the transfer function of our canonical micro-
circuit neural field model. Taking the Fourier transform of Equation (19) and sub-
stituting into Equation (17) gives:
T (k , ) ( 2 I 4 2i B B 2 J (k , )) 1 G
(20)
J (k , ) ABD(k , )
Substituting Equation (20) into Equation (15), we obtain an expression for the
predicted spectra as a mixture of contributions from each population weighted by
qa ( a 1,..., 4 ):
g ( , ) qaWa (k , )
a, k
2
(21)
Wa (k , ) e2 k
1me S a (k , ) R 1 (k , ) g u (k , )
2 2 2
cos(2ha k ), 2k sin(2ha k )
(22)
c caa i , c caa i
Qa (k , ) a 2 ( Daa(i ) (k , ) Daa
( e)
(k , )) a ma 2i a 2
(24)
Vab (k , ) Dab(i ) (k , ) Dba(i ) (k , ) 2 a b ma mb
In summary, the predicted spectral response at the sensor for the CMC field
model is given by:
g ( , ) e 2 1me a qa Sa (k , ) R 1 (k , ) g u (k , )
2
k
2 2 2
(25)
k
where the functions Sa (k , ) and R (k , ) are defined by (23). Equation (25) re-
flects the fact that the predicted spectral responses of the system are coupled to its
spatial as well as its temporal properties; these properties are encoded in the trans-
fer functions Sa (k , ) and R (k , ) through the underlying connectivity functions
Dab (k, ) . In turn, these are specified by the synaptic parameters associated with
the canonical microcircuit {mi , me , i , e , r ,} and the spatial parameters
{aab , cab , ha ,ab } that encode intrinsic and extrinsic connections among dif-
ferent layers and neighbouring columns or points on the cortical circuits.
The predicted spectral responses for the JR field model obeying Equation (8)
with connectivity determined by Equation (10) are also given by an equation of
the form of Equation (25) above (with a 1, 2,3 ) where the functions Sa (k , )
and R (k , ) are now given by:
D32( i ) (k , )
S1 (k , ) S 2 (k , ) ( e i) 2 ( i i ) 2
D31( i ) (k , )
S2 (k , ) D23(i ) (k , ) D31( i ) (k , ) 2 e i memi
S3 (k , ) D31(i ) (k , ) e me ( i i) 2
( i i ) 2 e 4 4i e 3 4i e 3 4 e 2 D13( i ) (k , ) D31(i ) (k , ) 2me 2 6 2
(26)
17
g y ( ) g ( , ) gn ( , ) y
u
gu ( , ) u , gn ( , ) n n , (27)
Re( ) ~ (0, ( , )), Im( ) ~ (0, ( , ))
and then turns to inferences on parameters, under the model selected. One usually
assumes the conditional density has a Gaussian form q( ) (, C) . This is
known as the Laplace assumption. The conditional density is quantified by the
most likely value of the parameters, and their conditional covariance C that
encodes uncertainty about the estimates and their conditional dependencies. Under
this assumption about the variational density and Gaussian observation noise, the
free energy has a very simple form:
( ) 12 ln | |
H 12 Re( ( ))T 1 Re( ( )) 12 Im( ( ))T 1 Im( ( ))
(28)
H 12 ( )T 1 ( ) 12 ln | | 12 ln | |
We will first formulate neural mass models as a special case of neural field
models by simply setting the conduction times to zero. This provides a useful per-
spective on the relationship between these two models, in terms of the implicit as-
sumptions we make when modelling observed data. A pragmatic advantage of
20
emulating neural mass models, with a transit time of zero, is that we can apply
precise shrinkage priors to conduction times to facilitate model comparison. In
other words, it provides a simple means of comparing models with and without
spatial dynamics (with and without prior constraints on conduction or transit
times). In particular, we first consider the mass and field variants of the JR model
and assume that output comes primarily from pyramidal cells. The corresponding
predictions for neural fields and masses are shown in Figure 4. These predictions
are based on model fits or inversions using local field potentials recorded from
primary (A1) auditory cortex in the Lister hooded rat, following the application of
the anaesthetic agent Isoflurane (see Moran et al., 2011 for details) under acoustic
white noise stimuli at a level of 83 dB. In brief, ten minutes of recordings were ex-
tracted from the continuous time domain data and down-sampled to a sampling
rate of 125 Hz. Frequency domain data-features were obtained from this epoch us-
ing a vector autoregression model of order eight. The model predictions of Fig. 4
illustrate nicely the difference between the field and mass models: one can see that
the neural field model has approximated the preponderance of low frequencies
more accurately than the neural mass model. This is because it has extra degrees
of freedom; namely conduction velocity and the extent of lateral connections.
These extend the repertoire of predictions to include those afforded by spatial dy-
namics. Crucially, the log-evidence for the neural field model was 1271 above the
log-evidence for the neural mass model. This suggests that there is a very strong
evidence for spatial dynamics over the cortical manifold in these auditory cortex
data (Penny et al., 2010). Recall that the model fit is based on optimising the free
energy bound to model-evidence. The free energy is just the difference between a
term quantifying accuracy (goodness of fit) and a term quantifying complexity.
This means the inversion provides explanations for empirical data that are both
accurate and parsimonious.
Figure 4: Real data (dashed line) and model predictions (full line) showing a 1 spectral profile that
is typically seen under anaesthesia. We observe a better fit of the field, relative to the mass model, in
particular for low frequencies.
Clearly, the choice of an appropriate model depends upon the question of inter-
est; in particular, neural fields are appropriate for addressing questions about the
deployment of sources on the cortical surface and induced spatial dynamics. How-
ever, the above example highlights that neural field models can be more appropri-
ate than mass models, from a Bayesian perspective, even if the spatial parameters
21
of a neuronal model are not the focus of study: In the context of our Bayesian
scheme, each model is scored using a free energy bound on model-evidence,
where better models have a higher free energy. This provides a principled way to
compare (score) different modes or hypotheses about how neuronal time-series are
generated.
We now turn to MEG data that summarise the spectral expression of endoge-
nous activity in the visual cortex of (twelve) human subjects described in detail in
Schwarzkopf et al., (2012). We used an adaptive spatial filter or beam-former
(Van Veen et al., 1997) to obtain estimates of ongoing neuronal activity in pri-
mary visual cortex. This provides estimates of electrical cortical activity based on
a weighted combination of sensors – sometimes referred to as a virtual electrode.
We then used the CMC model describing inter-laminar and lateral intra-laminar
interactions and inverted its mass and field variants. In these illustrative inver-
sions, the synaptic and spatial parameters were optimized and the intermodal dis-
tance ha was fixed to a physiologically plausible value. We computed the log evi-
dence ratio (using the free energy approximation) comparing field and mass
variants at the group level. As above, the neural mass model was formulated as a
special case of the neural field model by shrinking the conduction times to zero.
Contrary to our earlier result, we found no evidence in favour of the neural field
model (the relative log evidence between mass and field models was on average
2.62): see Fig. 5 for spectral fits of an exemplar subject.
Figure 5: Real data (dashed line) and model predictions (full line) for spectra in the gamma band ob-
tained from the human visual cortex during an attention task (Schwarzkopf et al., 2012). We observe
that the fits of both the field and mass models are equally good with no manifest differences.
This highlights the fact that the best model depends upon the data modelled. It
also underlines the importance of combining a neuronal model with a spatial for-
ward model: although both auditory and visual cortices are thought to conform to
22
the local homogeneity constraints implicit in neural field models, the loss of spa-
tial frequency resolution – with non-invasive data – might render neural field
models unnecessary, in relation to neural mass models. Our failure to establish a
significantly greater evidence for neural field models, in the present model com-
parison, is intuitively sensible because non-invasive MEG data have much lower
spatial resolution than the LFP data we used in the previous model comparison.
This observation speaks to the potential importance of using spatially resolved
data to take full advantage of neural field models. Data with high signal to noise
ratio and wide brain coverage – such as those afforded by ECoG sensors or multi-
array grids – can, in principle, disclose a full spectrum of spatiotemporal dynamics
at different scales, which may be important for an informed (efficient) estimate of
spatial parameters in neural field models.
Conclusions
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