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The major step
forward in
the modern theory of pattern formation was given by Turing
(1952), who used the linear analysis to determine the
conditions necessary for the creation of
spatial patterns in two-component reaction-diffusion
systems. A more recent criteria for pattern formation was
proposed by Koch & Meinhardt (1994) and Gierer &Meinhardt
(2000) and independently by Segel and Jackson (1972). They
postulate that the interplay between two antagonic feedbacks is
essential for pattern formation. On one hand,
the positive feedback consists in the
self-enhancement or autocatalysis of one of the
chemical components - generally called activator -, a
reaction necessary for small perturbations
to be amplified. On the
other hand, the increase in
activator's concentration must be complemented by a
fast-diffusing response in order to obtain pattern
formation.
The most
studied
examples of the two types of reaction-diffusion
systems are the Meinhardt system
(Gierer &Meinhardt 2000) and
the diffusive
Gray-Scott system (Pearson 1993) , respectively. The complex interplay
between activator and inhibitor or substrate chemical, aided by
the reaction and diffusion components
create most startling spatio-temporal patterns, such
as spots, stripes ,
travelling waves , spot
replication, and spatio-temporal chaos, in a
nutshell, a clear example of Turing patterns. The
Turing patterns are characterized by the active role that
diffusion plays in destabilizing the homogeneous steady state of the
system. They emerge spontaneously as the system is driven
into a state where it is unstable
towards the growth of finite-wavelength stationary
perturbations. Interesting enough, the replication
characteristic is a
particularity of the diffusive Gray-Scott
model alone, which
makes it the ideal model for
developmental research. In such
cases, cell-like localized structures grow, deform and make
replica
of themselves until they occupy the entire space .
The Turing patterns from the work of
Pearson 1993 on the
diffusive Gray-Scott model were confirmed
experimentally by
Lee, McCormick, Ouyang & Swinney (1993) including the spot
replication - Lee, McCormick, Swinney & Pearson (1994).
Theoretically, extensive work exist in the literature on the
dynamics
of this model concerning the ``spot replication'' in one, two
and
three dimensions (Muratov & Osipov 2000). Moreover, Muratov &
Osipov (1999) developed a theory of rotating spiral
waves for the
diffusive Gray-Scott model, as an example of a vivid
phenomenon
observed in many models and biological systems. In
addition,
Nishiura &Ueyama (1999) proposed a theoretical
mechanism that
drives the replication dynamics itself from a global bifurcation
point
of view.
This
model was originally introduced in Gray-Scott (1985) as an isothermal
system with chemical feedback in a continuously fed,
well-stirred tank reactor, where the last property implied the
lack of diffusion. The analysis of the system revealed stationary
states, sustained oscillations and even chaotic behavior. The model
considers the chemical reactions describing the
autocatalytic growth of an
activator V on the continuously fed substrate, U and
the decay of the former in the inert product P, subsequently
removed from the system. A major development was performed
by Pearson (1993) who introduced the role of space by
relaxing the constraint of a well-stirred tank and studied
the evolution of the concentrations of the
two chemical components, u(x,y,t) and v(x,y,t) in two
dimensions, in the limit of small diffusion. The system has as
parameters the Du and Dv, the
diffusion
coefficients, F, the dimensionless flow rate (the inverse of the
residence time) and k, the decay constant of the
activator, V. The original study involved
fixed diffusion coefficients, Du
= 2 x 10 -5 and Dv=10 -5,
with F and k
being the control parameters. |

For an overview of
the resultant patterns, one can integrate the equations
superimposing on the 2D space (X,Y) a gradient
of the control parameters, k and F. In such a way, one has
in each cell an approximate sample of the resultant pattern for the
associated F and k parameters. The
resultant ``phase
diagram'' is shown on the right, where besides spot
replication and stripes, the system shows for the bottom pairs of (F,k) travelling
waves and spatio-temporal chaos.
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As
discussed also in Pearson (1993), the spots occur only for
the parameter values for which the only steady state
is the red one - r.h.s. of the saddle-node
bifurcation curve --, and thus the gradient needed for the
formation and maintenance of the blue spots is an intrinsic
self-sustaining feature of the system. Once a spot of
high V is formed, it is maintained by the concentration
difference between the its center and the
surroundings of the spot. As the concentrations are
limited to the [0,1] interval, the spot can grow until
its maximal gradient is not enough
to achieve a maximum concentration in
the center and thus its
V-value starts to decrease. This induces the spot-division
phase.
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Mazin
et al.(1996) account for and illustrate the Turing space - the region
in the parameter space for which the blue state is unstable with
respect to the growth of standing spatial perturbations. Their
analysis clearly determines the crucial role played
by the ratio of the diffusivity coefficients in the pattern
formation. The increase of the diffusivity ratio beyond 2 ,
the value investigated in the original paper of
Pearson(1993), leads to a significant extension of the Turing
regime.
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Lesmes et al.
(2003) have carried out the first study of the
noise-controlled pattern formation in the Pearson model, with
emphasis
on the self-replicating patterns. They found that for the chosen value
of the pair (F,k), the
noise drives the system from the
non-multiplicative, stripe-like pattern to the
spot-multiplication
one .
Interesting enough, they argue in favor of an optimal noise
intensity, A for which the
number of spots is maximal, after a
sufficiently long integration time .
In comparison with the results of Lesmes et
al., ours
suggest the existence of an
interval of optimum noise-intensity values leading to
a maximum number of spots, rather than a single
optimum value. In other words, our distribution
of spots shows a plateau at the level of maximum number of
spots, while theirs appears to have
a more bell-like shape. The
plateau characteristics is more evident
in the distribution of the area occupied
by the two features -- spots and lines
- as the noise intensity varies.
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Recently, Robert Munafo drew my attention to additional interesting studies of this system.
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